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Volume 32 1978 Number 1

JOURNAL

of the

LEPIDOPTERISTS’ SOCIETY

Published quarterly by THE LEPIDOPTERISTS’ SOCIETY

Publié par LA SOCIETE DES LEPIDOPTERISTES
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN
Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS

7 April 1978

THE LEPIDOPTERISTS’ SOCIETY

EXECUTIVE COUNCIL

J. W. Trpen, President KENELM W. Putuip, Vice President
I. F. B. Common, Ist Vice President JuLian P. DoNAHuE, Secretary
Lionet Hiccrns, Vice President RONALD LEUSCHNER, Treasurer

Members at large:

F. S. CHEW R. A. ARNOLD J. F. EMMEL
D. F. Harpwick E. D. CAsHATT R. R. GATRELLE
J. B. ZrEcLER R. E. STANFORD A, P.\ Prats

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formally constituted in December, 1950, is “to promote the science of lepidopterology
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Memoirs of the Lepidopterists’ Society, No. 1 (Feb. 1964)
A SYNONYMIC LIST OF THE NEARCTIC RHOPALOCERA

by Cyrit F. pos Passos

Price: Society members, $5.00 U.S.; non-members, $7.50 U.S. Paper covers, revisions
of the Melitaeinae and Lycaenidae supplied separately.

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Cover illustration: Dasychira dorsipennata larva, dorsal and lateral views. From
Fascicle 22.2, “Lymantriidae,” by Douglas C. Ferguson, in Moths of America North
of Mexico. The drawing was done by E. R. Hodges, Scientific Illustrator, Department
of Entomology, Smithsonian Institution. (Reproduced by permission of the author.)

JOURNAL OF

Tue LEpPIDOPTERISTS’ SOCIETY

Volume 32 1978 Number 1

Journal of the Lepidopterists’ Society
32(1), 1978, 1-2

ieee OF PARNASSIUS CLODIUS GALLATINUS
(PAPILIONIDAE )

STEVE KOHLER

Montana Department of Natural Resources and Conservation, Division of Forestry,
2705 Spurgin Road, Missoula, Montana 59801

ABSTRACT. Types of Parnassius clodius gallatinus Stichel, 1907, were discov-
ered in the remnants of the Elrod collection, at the University of Montana, Missoula.
Proper labels have been attached to the specimens, and they have been placed in
the collection of the American Museum of Natural History.

The description of Parnassius clodius gallatinus Stichel was based on
a pair illustrated by Elrod (1906). The actual specimens were never
seen by Stichel (1907).

While preparing a paper clarifying the nomenclature of Parnassius
clodius Ménétriés subspecies found in the Rocky Mountains (Ferris,
1976), Cliff Ferris contacted me in an attempt to locate the types of
gallatinus. A search through the remnants of the Elrod Collection,
housed at the University of Montana, Missoula, led to the discovery of
the two specimens matching El!lrod’s 1906 illustrations. They were
figured on page 16 of “The Butterflies of Montana.”

At the suggestion of Ferris, the two specimens have been placed in
the collection of the American Museum of Natural History. The follow-
ing labels are affixed to the specimen pins:

Holotype Male: A white label partially machine printed in black ink
and partially hand lettered in red ink which reads: Gallatin Co. Mont./
Eley. 6800/ Col. E. Koch/ 6-27 1900, and a red label hand lettered in
black ink which reads: Holotype ¢/ Parnassius clodius/ gallatinus
Stichel/ ex. Elrod Coll. Univ./ Mont. S. Kohler 1976.

Allotype Female: A white label partially machine printed in black ink
and partially hand lettered in red ink which reads: Gallatin Co. Mont./
Elev. 6800/ Col. Cooley/ 6-27 1900, and a red label hand lettered in

bo

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

te

Fig. 1. Parnassius clodius gallatinus Stichel: a) holotype male, dorsal; b) same,

ventral; c) allotype female, dorsal; d) same, ventral. Photos approximately 7
natural size.

black ink which reads: Allotype ?/ Parnassius clodius/ gallatinus Stichel/
ex. Elrod Coll. Univ./ Mont. S. Kohler 1976.

The specimens are in good condition except for some minor dermestid
damage to the abdomen of the female. An additional pair collected by
Cooley at the same locality and on the same date was located in the
collection of Montana State University, Bozeman, for Ferris by Dr. Nor-
man L. Anderson. These are not designated as paratypes.

Because of the scarcity of copies of Elrod’s “The Butterflies of Mon-
tana, the type specimens have been illustrated in Figure 1.

ACKNOWLEDGMENTS

I would like to thank Dr. James H. Lowe and the University of Mon-
tana for allowing the type specimens to be placed at the American
Museum of Natural History, and also Cliff Ferris for comments and
assistance,

LITERATURE CITED

IeLnop, M. J. 1906. The butterflies of Montana. Univ. Montana Bull. 10: 1-174.

Fennis, C. D. 1976. A note on the subspecies of Parnassius clodius Ménétriés
found in the Rocky Mountains of the United States (Papilionidae). J. Res.
Lep. 15: 65-74.

SticHEL, H. 1907. Lepidoptera Rhopalocera Fam. Papilionidae Subfam. Parnas-
siinae. Wytsman Gen. Ins. Fasc. 58: 1-60.

Journal of the Lepidopterists’ Society
32(1), 1978, 3-19

SPECIFICITY, GEOGRAPHIC DISTRIBUTIONS, AND
FOODPLANT DIVERSITY IN FOUR CALLOPHRYS
(MITOURA) (LYCAENIDAE)

Kurt JOHNSON

Department of Biology, City University of New York, City College, Convent
Avenue and 138th Street, New York, New York 10031

ABSTRACT. The species C. siva, gryneus, hesseli, and turkingtoni are examined.
Genitalic evidence of their non-conspecificity is provided along with discussion of
particular localities of sympatry. Detailed distributional data are illustrated and a
documented table of foodplant diversity included. C. siva and gryneus are oligoph-
agous On numerous species of Juniperus (Cupressaceae) which replace each other
geographically across the United States. C. hesseli is monophagous on Chamaecyparis
thyoides (Cupressaceae); the foodplant of turkingtoni is unknown. Evidence indicates
that all local populations are specific to one foodplant species.

Callophrys ( Witoura) nelsoni ( Boisduval), C. siva (Edwards), C. loki
(Skinner), C. gryneus (Hiibner), and C. hesseli (Rawson & Ziegler),
aside from taxonomic descriptions, have been subject to several biological
and regional studies, but published works (Anderson, 1974; Johnson,
1972; Pease, 1963; Rawson et al., 1951; Remington & Pease, 1955) are
very heterogeneous in content and comprehensiveness.

During the last four years I have been compiling data on their dis-
tributions and larval foodplants as a base for taxonomic studies of the
group. I have also been studying the genitalia of all Nearctic and
Neotropical Callophrys (Mitoura) in detail (Johnson, 1976a). The pur-
pose of this paper is to present detailed distributional data for three of
these species (C. siva, C. gryneus, and C. hesseli), demonstrate that C.
siva and C. gryneus are not conspecific, and summarize data on larval
foodplants, a number of which are new to the literature. What bio-
geographical data are known on the newly named C. turkingtoni Johnson
(Johnson, 1976b) will also be presented. The specificity of C. siva and
C. nelsoni involves several complex problems in the northwestern United
States and will be treated in a separate paper (Johnson, 1977).

METHODS AND MATERIALS

Using the collection of the American Museum of Natural History as a
basis, additional information on localities and possible local foodplants
was gathered by correspondence and recorded county by county. Speci-
mens or photographs were solicited in cases of peripheral or isolated
populations, and available published records were included. The re-
search aimed at definitive treatment on the species level only. Genitalic

+ JOURNAL OF THE LEPIDOPTERISTS SOCIETY

studies of males and females were performed in areas where C. siva and
C. gryneus were reportedly sympatric. These genitalia were compared
with those from many parts of the ranges of C. siva and C. gryneus, as
well as with dissections of other congeners. The number of these speci-
men dissections included: C. siva, 78; C. gryneus, 46; C. hesseli, 14; C.
turkingtoni, 1; C. nelsoni, 83; C. rosneri, 46 (Johnson, 1976a); C. barryji,
19 (Johnson, 1976a); C. byrnei, 9 (Johnson, 1976a), and C. loki, 15.
Geographic ranges were studied to discover areas of insect distribution
not coinciding with present published foodplant knowledge, and efforts
were then made to make the list for each species complete by identifica-
tion of exact plants with which the adults were associated by perching
behavior (Johnson & Borgo, 1976) or on which oviposition or larvae were
observed. Full documentation of each of these methods is given in the
foodplant table (Table 1) since a degree of fallibility has been demon-
strated in each (Brower, 1958; Downey & Dunn, 1965). An ongoing
effort to compile foodplant specimens at one institution was initiated,
and plants collected thus far are cited in the table. Since the perching
behavior of these insects limits general flight patterns to the vicinity of
the foodplant, and since data not only in this study but another (John-
sonn, in prep. ) indicate that C. siva and C. gryneus are exclusive Juniperus-
feeders, some useful evidence on larval foodplants in areas where only
one juniper species was regionally present could be culled from identifi-
cation of the plants at the locality indicated on the specimen labels.
The list of plants established as the only Juniperus species present in a

region (R) or at a locality (L), source of butterfly data (B), source of
plant data (P) is:

C. siva: Juniperus deppeana Steud., (LL) 10 mi. NW Pine Springs, Culberson
Co., Texas, (B) R. O. Kendall, (P) Herbarium, University of Texas, Austin; J.
deppeana, (1.) 5 mi. W of McDonald Observatory, Jeff Davis Co., Texas (Bk. 'O:
Kendall, (P) Herbarium, University of Texas, Austin. Juniperus occidentalis oc-
cidentalis Hook. x J. osteosperma Torr. (Little), (R) Washoe Co., Nevada (Reno
and vicinity westward), Ormsby and Douglas cos., (B) P. Herlan, (P) Vasek,
1966. Juniperus monosperma (Engelm.) Sarg., (LL) Sycamore Canyon, NW of
Nogales, Santa Cruz Co., Arizona, (B) Share and Clark (American Museum of
Natural History (AMNH)), (P) Herbarium, Arizona State University, Tempe.
Juniperus pinchotii Sudw., (R) Reeves Co., Texas, (B) D. Stallings and M. R.
Turner, (P) Adams, 1972, R. P. Adams, pers. comm. C. gryneus: Juniperus
virginiana L., (R) Cass Co., Texas, (B) R. O. Kendall, (P) Adams & Turner,
1970, R. P. Adams, pers. comm. Juniperus ashei Buchholz, (R) McLennan Co.,
Texas, (B) R. O. Kendall, (P) Adams, 1972; Adams & Turner, 1970; R. P. Adams,
pers. comm. Juniperus pinchotii Sudw., (R) Pecos Co., Texas, (B) R. O. Kendall,
(P) Adams, 1972, R. P. Adams, pers. comm. Juniperus deppeana Steud., (L)
Huejotitlan, Chihuahua, Mexico, (B) AMNH, (P) Little, 1971. J. deppeana, (i)
Baboquivari Mountains, S of Baboquivari Peak, Pima iGo Arizona,” (ByenieaD:
Gunder (AMNH), (P) Herbarium, Arizona State University, Tempe. Juniperus
virginiana L. * J. horizontalis Moe nch., (1) Lynxville, along Mississippi River,

VoLUME 32, NUMBER 1

Aste I.

Larval foodplants established by the identification of exact plants.

Foodplant taxa and specimens

Butterfly taxa and specimens

Callophrys (Mitoura) siva

Juniperus scopulorum Sarg."

Plants, PL: Van Haverbeke, N-2;
hybrid index at site—63 + 8% J.
scopulorum; SL: Johnson, 1972.”

Plant specimens, K. Johnson #2 (Smiley
Canyon = VanH. N-2; #1 (Chadron),
Royal Ontario Museum, Toronto

(ROM ).°

Juniperus scopulorum var. columnaris
Fasset

Plants, PL: Van Haverbeke ND-S, I: T.
McCabe, (H) North Dakota State
Univ., Fargo; SL: Van Haverbeke, 1968.
Plant specimens, Van Haverbeke ND-8,
Univ. of Nebraska (UN).

Juniperus scopulorum Sarg. X J. virgini-
ana L.

Plants, PL: Van Haverbeke N-7;

hybrid index at site—48 + 6% J.
scopulorum; Range of use of hybrids
(Johnson, 1972 )—70 + 4% to 36 + 4%
J. scopulorum.

Plant specimens, Van Haverbeke N-7,
UN; K. Johnson #3 (Sizer, Keith Co. )
ROM.

Juniperus virginiana L.

Plants, 17 mi S of PL: Van Haverbeke
N-4; hybrid index at site—36 + 4%

J. scopulorum; SL: Johnson, 1972.
Plant specimens, Van Haverbeke N-4,
UN; K. Johnson #4 (locality as above )
ROM.

Prostrate morph of J. scopulorum Sarg.
x J. virginiana L. x J. horizontalis
Moench.

s. siva; Dawes Co., Nebraska (Smiley
Canyon ), W of Fort Robinson; Catholic
Cemetery, Chadron.*

Butterthes, PE (ACS@: B)*: K.
Johnson (AMNH).°

s. siva; Slope Co., North Dakota
(Amidon, along burning coal vein).
Butterflies, PC: T. McCabe.

s. siva; Garden Co., Nebraska (bluffs
above N. Platte River, nr. Lewellen).
Butterflies, PC (AC): L. Running,
AMNH.

s. siva; Rock Co., Nebraska (Long Pine
Rec. Area).

Butterflies, PC (AC, LC, B): K.
Johnson, L. Running, AMNH.

s. siva; Saskatchewan, Canada (Val
Marie, near Rosefield along Frenchman
River ).

pm
1 Taxon of foodplant used (according to nomenclature of USDA (1953) and Little (1971)).

2Source of plant data:

PL — butterflies were specifically collected at a_ particular locality

studied by Cupressaceae taxonomists; their designation of the site is noted along with the date
of their study. “Hybrid index” refers to these studies’ calculation of the degree of hybridity in
plants at these areas. Plant identifications are noted as “7”: I, exact substrate plant identified
pen Ee —Rerbarum at ==»), and! location of voucher specimens; I,, plant identified

by data sent to ______ by

; I,, foodplant established in original description of butterfly,

citation given; I,, plant identified from specimens sent to ——_——_; I,, substrate plant established

by matching herbarium specimens with butterfly data and establishing that no other species co-
occurs, herbarium cited. An “*” following this category (‘‘Plants’”’) means this foodplant usage

is well

3 The label number and place of deposition of plant specimens collected in this study An “*
means collection in progress at time of this writing.

known; SL = other literature which supports this identification.

>?

4Taxon of butterfly concerned (as designated in annotated list), with state and exact locality.

5 Source of butterfly data: PC = “personally collected by

Letters in parentheses

following mean: AC, adults commonly observed perching; AI, adults perching but not commonly
observed; LC, larvae collected; LO, larvae observed; B, behavior studied in detail; O, oviposition

observed.

6 Location of specimens if not aforementioned person (AMNH =the American Museum of
Natural History, New York). M =museum specimens were used as the source of data; ver.

means verified by

, and method. TL = type locality of the insect.

6 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TAsrE

Continued

Foodplant taxa and specimens

Plants, Schurtz (1971) indicates this
area would be included in his tri-
parental swarm. Van Haverbeke (pers.
comm.) supports this evaluation; I: (H)

Univ. Saskatchewan, Regina; I: D. F. Van

Haverbeke (data from R. Hooper, K.
Johnson); SL: Little, 1971; Van
Haverbeke, 1968; Fassett, 1945.

Plant specimens, (H) Univ. Sask. (Sask.
Prairie Park); K. Johnson #5* ( Hooper )
ROM.

Juniperus osteosperma ( Torr.) Little
Plants, I;: D. F. Van Haverbeke; SL:
Emmel & Emmel, 1973; Johnson, 1977;
Little, 1971.

Plant specimens, K. Johnson #15
(Running, locality as above) AMNH.

Juniperus californica Carr.
Plants +, I: D. F. Van Haverbeke; SL:

Comstock, 1927; Emmel & Emmel, 1973.

Plant specimens, K. Johnson #16
(Leone, locality as above) AMNH.

Juniperus occidentalis occidentalis Hook.
Plants, I: D. F. Van Haverbeke; SL:
Johnson, 1977, Little, 1971.

Plant specimens, K. Johnson #17
(Buckingham, locality as above).

Juniperus occidentalis australis Vasek
Plants, I: John H. Lane; SL: Vasek,
1966.

Butterfly taxa and specimens

Butterflies, PC (AC): R. Hooper (ver.
K. Johnson, photo ).

s. ssp.; White Pine Co., Nevada (nr.
McGill Junction).

Butterflies, PC (AC): L. Running,
AMNH.

s. juniperaria; Los Angeles Co., Cali-
fornia (Mint Canyon).

Butterflies, PC (AC): M. Leone,
AMNH.

s. ssp.; Jefferson Co., Oregon (nr. Warm
Springs, W on road to Twin Buttes ).
Butterflies, PC (AC): F. Buckingham,
AMNH.

s. ssp.; Tulare Co., California (vic.
Kennedy Meadows ), San Bernardino
Co., California (Big Bear Lake).
Butterflies, PC (AC): John H. Lane.

Callophrys (Mitoura) gryneus

Juniperus virginiana L.

Plants, I: K. Johnson; SL: Klots, 1951,
Little, 1971.

Plant specimens, K. Johnson #21
(locality as above).

Juniperus silicicola (Small) Bailey
Plants, I: F. D. Fee, (H) Univ. Florida,
Gainesville; L: (H) Univ. Gainesville;
SL: Klots, 1951; Little, 1971.

Plant specimens, K. Johnson #10 (St.
Augustine locality, Univ. Florida) ROM.

Juniperus scopulorum Sarg. * J. virgini-
ana LL.

Plants, PL: Van Haverbeke M-1; hybrid
index near site—27 + 4% J. scopulorum;
Range of use of hybrids (Johnson,

g. gryneus; Ulster Co., New York ( West
Park, Holy Cross Publications ).
Butterflies, PC (AI): K. Johnson,
AMNH.

g. sweadneri; St. Johns Co., Florida
(along Ocean Rt. A1A, St. Augustine ).
Butterflies, PC (AC): F. D. Fee.

g. gryneus; Jackson Co., Missouri
(general ).
Butterflies, PC (AC): J. R. Heitzman.

VoLUME 32, NuMBER 1

TABLE l.

Continued

Foodplant taxa and specimens

Butterfly taxa and specimens

1972 )—38 + 4% J. scopulorum to
27 + 4% J. scopulorum.

Plant specimens, K. Johnson #11
(Heitzman, Independence ) ROM.

Juniperus ashei Buchholz

Plants, I: J. R. Heitzman; SL: Little,
1971.

Plant specimens, K. Johnson #12*
(Heitzman, ?) ROM.

Juniperus pinchotii Sudw.
Plants [two examples], L: R. P. Adams
(Scott, Roever); SL: Adams, 1972.

I: R. O. Kendall; SL: Little, 1971;
Adams & Turner, 1970.

Juniperus virginiana L. x J. horizontalis
Moench.

Plants, I: (by reason of Schurtz, 1971)
D. F. Van Haverbeke; SL: Schurtz,
1971; Little, 1971.

Plant specimens, K. Johnson #13
(locality as above).

g. ssp.; Barry Co., Missouri (Eagle
Rock ), also McDonald Co.; Washington
and Carroll cos., Arkansas.

Butterflies, PC (AC): J. R. Heitzman.

g. castalis [two examples]; Armstrong
Co., Texas (just below N rim of Palo
Duro Canyon, 15-16 mi. S Claude).

Butterflies, PC (AC): M. Toliver,

H. A. Freeman, J. M. Burns, K. Roever
(R. O. Kendall); J. Scott.

Bexar Co., Texas (Reo Seco Road, off
U.S. Hwy. 281 N of San Antonio).
Butterflies, PC (AC, O): R. O. Kendall.

g. gryneus; Dane Co., Wisconsin (10
mi. W of Madison).
Butterflies, PC (AC): W. Sieker.

Callophrys ( Mitoura) hesseli

Chaemaecyparis thyoides (L.) B.S.P.
Plants*, I: S. Hessel, G. W. Rawson,

J. B. Ziegler; I,: Rawson et al., 1952;
Rawson & Ziegler, 1950 (therein det. by
I. M. Johnston, Harvard Univ. ).

hesseli; Ocean Co., New Jersey
(Lakehurst, TL).

Butteniess 2G. ( ACO. LEG) S.
Hessel, G. W. Rawson, J. B. Ziegler.

Lacrosse Co., Wisconsin and 5 mi. W of Sauk City, Sauk Co., Wisconsin, (B)
F. Amold and W. E. Sieker, (P) Ross & Duncan, 1949; Schurtz, 1971; D. F.
Van Haverbeke, pers. comm. C. turkingtoni: Juniperus flaccida Schlecht., (R)
10 mi. E of Namiquipa, Chihauhua, Mexico, (B) W. Gertsch and M. Cazier
(AMNH), (P) Little, 1971; Herbarium, University of Mexico, Mexico City.

RESULTS
Genitalia of C. gryneus and C. siva
Genitalia of males and females were studied in three regions where
these species were reportedly sympatric (Davis Mountain, Texas; Guada-

lupe Mountains, New Mexico and Texas; and Baboquivari Mountains,
Arizona) and found to be easily separable. However, some traditionally

8 JoURNAL OF THE LEPIDOPTERISTS SOCIETY

Figs. 1-9. Female genitalia of selected Nearctic Callophrys (Mitoura) spp.
C. siva siva: 1, topotypical; 2, heavily sclerotized areas of lamellae and eighth
sternite; 3, showing tufts of “hair” ailowing diagnosis by naked eye. C. gryneus:
4, topotypical. C. hesseli: 5, topotypical. Genital plates of sympatric species near
Alpine, Texas: 6, C. gryneus and 7-9, C. siva.

used wing-pattern characters for distinguishing these species (ventral
secondaries: post basal spots or pattern of mesial band) were shown to
be less reliable (also noted in Johnson, 1976a, 1977). The diagnostic

genitalic characters are as follows:

Females (Figs. 1-9). C. gryneus (Figs. 4, 6): ductus bursa longer and not
“club-ended” as on siva; lamellae tapering caudad from antrum, not shouldered
as on siva, lamellae postvaginalis nearly as long as broad; juncture of lamellae
and eighth abdominal sternite not heavily sclerotized or connected,

C. siva (Figs. 1-3, 7-9). Ductus bursa shorter than gryneus and “club-ended”;
lamellae distinctly shouldered, lamella postvaginalis much broader than long. Junc-
ture of lamellae and eighth abdominal sternite heavily sclerotized, in area between

VOLUME 32, NUMBER 1 9

Figs. 10-12. Male genitalia of selected Nearctic Callophrys (Mitoura) spp.,
lateral and posterior views with tip of aedeagus (right) and falces (left): 10, C.
gryneus castalis, topotypical; 11, C. siva siva, topotypical; and 12, C. hesseli, topo-
typical.

1. postvaginalis and 1. antevaginalis forming bulkly ridges and convolutions at their
juncture, these binding lamellae tightly with eighth abdominal sternite and con-
taining many spines.

[C. hesseli (Fig. 5). Easily recognized by unique shape of the lamellae and
broad cephalad tapering from the antrum (figured for reference). C. turkingtoni:
female unknown. ]

Males (Figs. 10-12). C. gryneus (Fig. 10). Valvae, lateral shape: only barely
concave between dorsal and ventral articulation with vinculum; valvae, caudad
saccus (dorsal or ventral view): rounded and indented, vaguely shouldered caudad.
Saccus: long and broad.

C. siva (Fig. 11). Valvae, lateral shape: deeply concave and rounded between
dorsal and ventral articulation with vinculum; valvae, caudad saccus (dorsal and
ventral view): parabolic and unindented, no shouldering caudad. Saccus: short
and much less broad than gryneus.

[C. hesseli (Fig. 12). Lateral shape of valvae less broad, quite concave between
articulations with vinculum, and much longer caudad; valvae caudad saccus broadly
round, indented, and extremely shouldered caudad (figured for reference). C.
turkingtoni (Johnson, 1976a), easily recognized by extremely long caudad extension
of valvae and by heavily sclerotized and spiny area of valvae, caudad saccus.]

JouRNAL OF THE LEPIDOPTERISTS SOCIETY

10

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VoLUME 32, NuMBER 1 ai

& —
J. virginiana =

v
6)
gs J. flaccida
ZN J. californica

= J. pinchotii

poles occidentalis
ee communis

horizontalis

BY J. scopulorum

Se
J. monosperma

fi J. deppeana
EE

Eo J. osteosperma
FA) preceding 4

ia J. ashei

y ee 7 gryneus | I &
¢ Se YW a &
C. (M.) siva siva JT Known transplanted (x) ¢- (M.) siva juniperaria
@ green morph Bop aeton: Oc. (M.) siva mansfieldi
@ brown morph 2? ee OH CEE (Oc. (M.) siva, inner coastal
relationship with Danpenbaoeninoenh
green-brown Cc. (M.) nelsoni complex. z ae

we Cc. (M.) siva, high altitude
morph
(M.) turkingtoni

contact zone

Sympatric C. (M.) siva and
alc. (M.) gryneus, diagnosed
by genitalia. @c.

Mexican populations assigned to species as indicated.

Fig. 14. Nearctic distributions of Callophrys (Mitoura) siva and its larval food-
plant Juniperus spp., and known range of C. (M.) turkingtoni. Plant distributions
adapted from Little, 1971. Distribution of Juniperus horizontalis shown only as it
exceeds J. scopulorum northward; hybrid swarms of Juniperus spp. illustrated in

Pig, 1d.

12 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

ae re ew were ee ene pent es tors an

North Dakota

r South Dakota

Wyoming
Nebraska

Mig. 15. Bi-parental and tri-parental swarms of divergence in Nearctic Juniperus:
top, localities indicated by Van Haverbeke (1968) as “hybrid” J. virginiana x J.
scopulorum; center, parental areas of J. horizontalis indicating possible centers for

VoLUME 32, NuMBER 1 13

Geographic Distributions and Available Larval Foodplants

Figs. 13 and 14 show the Nearctic distributions of species of Callophrys
(Mitoura) in relation to the ranges of available or established larval
foodplants. Fig. 15 shows areas of Juniperus ranges that have been
botanically demonstrated as “hybrid swarms” (Van Haverbeke, 1968;
Schurtz, 1971).

Summary of Data and Current Taxonomic Usages

The following is a review of the current common usage of trinomens in
each group with a summary that includes distribution, foodplant(s) as
established in this paper, general comments on the phenotype, and notes
on the particular significance of each population. Where populations are
under study by other lepidopterists, and especially where they are plan-
ning to assign new names, | have called these subspecies “ssp.” and in-
cluded the appropriate investigator's name in brackets.

Annotated List
Callophrys (Mitoura) siva

C. (M.) siva siva. Type locality: Fort Wingate, McKinley Co., New Mexico.
Distribution: Workers have named populations distinct from this taxon only on
the West Coast, although others are undoubtedly present. Phenotype: There are
two general morphs, based on ground color of the ventral secondaries. Great Basin
populations (Fig. 14, squares) are brown beneath, whereas others (Fig. 14, plain
black circles) are green. Populations of green-browns and mixed greens and browns
occur in western Utah (Fig. 14, overlapping square and circles). The brown morph,
which Peter Herlan, H. K. Clench, and I have investigated (Johnson, in prep.),
is separately treated below. Foodplants: Many western Juniperus species (see
Table 1) replace each other geographically. “Hybrids” (see Summary and Con-
clusions) of J. virginiana x J. scopulorum, J. virginiana x J. horizontalis, and J.
virginiana X J. scopulorum x J. horizontalis (possibly also J. occidentalis x J. osteo-
sperma) occur in western Nevada, adjacent California, and eastern Oregon. All
populations of Callophrys siva are on erect trees except for one local population
(Val Marie, near Rosefield, Saskatchewan, along Frenchman River) on prostrate
plants.

C. (M.) siva ssp. [Johnson, in prep.]. The research of Herlan, Clench, and
Johnson involves naming this Great Basin population. Distribution: southern
Nevada northward to Idaho; brown eastward to Salt Lake City; brown westward
to southeast Oregon; broad interface with green morph C. (M.) siva siva in western
Utah (e.g., Eureka, Dividend, Provo, western Millard Co.). Foodplants: Herlan
reported J. osteosperma, but probably J. osteosperma x J. occidentalis near Reno,
Nevada (Vasek, 1966). Apparently not J. scopulorum where it is available, al-

<<

“hybrid” status with J. virginiana; bottom, localities indicated by Schurtz (1970)
as “hybrid” J. virginiana <x J. scopulorum xX _ J. horizontalis.

14 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

though Idaho occurrences of C. (M.) siva may be lacking due to inadequate
sampling.

C. (M.) siva juniperaria (Comstock). Type locality: Los Angeles Co., California.
Distribution: western San Bernardino, northern Los Angeles, eastern Kern, and
southern and northern Inyo cos.; transition with next subspecies in northern Ventura
and northeastem Santa Barbara cos. Foodplant: Juniperus californica throughout
its range, but in one locality (areas between Phelan and the San Gabriel Mountains,
close to the Los Angeles, San Bernardino county lines) it is known to perch also on
J. osteosperma (J. Lane, pers. comm. ).

C. (M.) siva mansfieldi (Tilden). Type locality: Simmler, San Luis Obispo
Co., California. Distribution: southern California—western Kern and _ eastern
San Luis Obispo cos. This is somewhat northward of the preceding subspecies.
This taxon is ill-defined from that immediately below. Phenotype: deep green
morph. Foodplant: J. californica.

C. (M.) siva ssp. [Lane, A]. Distribution: north of the preceding entity in the
south inner coastal range of California (northern San Luis Obispo, southwestern
Fresno, San Benito, eastern Santa Clara, and western Contra Costa cos.). Pheno-
type: brown morph. Foodplant: J. californica.

C. (M.) siva ssp. [Johnson, in prep.]. The identity cf reputed C. (M.) siva
specimens from eastern Oregon and Washington to western Idaho, and their rela-
tion to the name C. (M.) nelsoni, has been resolved by detailed genitalic studies
(Johnson, in prep.). Populations in nearly all of Oregon east of the Cascades to
extreme southwest Idaho and extreme southeastern Washington are C. siva. Pheno-
type: burgundy-brown morph. Foodplants: Oregon, Washington—J. occidentalis;
Idaho—J. osteosperma. The taxon in press by Johnson includes only the J. occi-
dentalis utilizers; those on J. osteosperma represent northward range of another
subspecies distributed throughout Nevada and reviewed above.

C. (M.) siva ssp. [Lane, B]. Distribution: High altitudes in the southern Sierra
Mountains (e.g., Kennedy Meadows, Tulare Co., California) and San Bernardino
Mountains (e.g., Big Bear Lake, San Bernardino Co., California). Phenotype:
green morph. Foodplant: J. occidentalis australis.

Callophrys (Mitoura) gryneus

C. (M.) gryneus gryneus. Type locality: Rappahanock Co., Virginia. Distribution:
eastern and central North America in scattered populations wherever J. virginiana
occurs. Phenotype: green morph. Foodplants: J. virginiana, but not observed
feeding on sympatric J. communis, prostrate morphs of J. horizontalis x J. virginiana,
or prostrate morphs of J. horizontalis northward; however, apparently utilizes erect
morphs of J. horizontalis > J. virginiana (see Summary and Conclusions). Particular
note: sometimes collected on nectar sources with C. (M.) hesseli, but foodplants
are segregated by habitat in nature and not interchangeable.

C. (M.) gryneus sweadneri (Chermock). Type locality: St. Augustine, St. John’s
Co., Florida. Distribution: Florida, perhaps southern Georgia, and north along
the Atlantic Coast where J. silicicola occurs. Phenotype: green morph. Foodplant:
J. silicicola.

C. (M.) gryneus castalis (Edwards). Type locality: McLennan Co., Texas.
Distribution: mainly Texas, but also Chihuahua, Mexico, and areas west of the
Mississippi River “gap” in juniper ranges; in addition, used by some workers as a
form name within eastern United States populations. Phenotype: green morph.
Foodplants: J. virginiana, J. ashei, and J. pinchotii, replacing each other westward.
J. deppeana and possibly J. flaccida in Mexico.

C. (M.) gryneus ssp. [Johnson, in prep.]. Distribution: the Baboquivari Moun-
tains eastward into Cochise Co., Arizona, and possibly southward in disjunct ranges
of J. deppeana. Phenotype: green morph. Foodplant: J. deppeana suspected.

VoLUME 32, NuMBER 1 15

Callophrys (Mitoura) hesseli

C. (M.) hesseli. Distribution: see Fig. 13. Phenotype: green morph. Food-
plant: Chamaecyparis thyoides.

Callophrys (Mitoura) turkingtoni

C. (M.) turkingtoni, a single specimen known from Namiquipa, Chihuahua,
Mexico, in habitat of J. flaccida. Phenotype: brown morph.

SUMMARY AND CONCLUSIONS

Interspecific relations. Studies of C. siva and C. gryneus at several
sympatric localities (21 specimens from the Baboquivari Mountains,
Pima Co., Arizona; Cochise County (general), Arizona; Guadalupe
Mountains, Eddy and Otero cos., New Mexico, Culberson Co., Texas;
and Alpine, Brewster Co., Texas) confirmed that they are separable by
genitalia of the males and especially the females (Johnson, in prep.).
Since town, county, or mountain range is the only data available on some
of these specimens, the extent of their microallopatry or microsympatry
remains unknown. Biogeographic data suggest that the species may be
altitudinally separated at some localities in Texas (C. siva on higher
altitude J. deppeana, C. gryneus on lower altitude J. pinchotii), but it is
likely that interspecific competition occurs at some locations. Sharing of
nectar sources may occur, as reported in C. gryneus and C. hesseli (J. B.
Ziegler, pers. comm.). These two species are generally segregated by the
habitats of their foodplants. The female genitalia of C. hesseli have not
been previously figured in the literature and are included in Fig. 5.

Foodplant relations. C. siva and C. gryneus utilize a broad spectrum
of related and equally acceptable Juniperus species, which replace or
exceed each other in geographic distribution over the Nearctic Realm.
There is evidence that every species of Juniperus in the Nearctic is
utilized, with two exceptions: J. communis L. and J. horizontalis Moench.
Van Haverbeke (1968) and especially Schurtz (1971) have shown that
J. horizontalis is actually part of a broadly distributed “swarm of diver-
gence’ which involves the parental stock to which the names J. virginiana,
J. scopulorum, and J. horizontalis have been applied. Van Haverbeke
(pers. comm.) prefers Schurtz’s interpretation that each of these merits
species status but that they are tied by their evolutionary histories, J.
virginiana being an eastward evolutionary manifestation of J. scopulorum
and J. horizontalis being a northward evolutionary manifestation of this
biparental parent stock. Thus, there is little chemical or morphological
reason (unless it is the number of needles versus fleshy leaves) that
would prevent use of J. horizontalis by these Callophrys (Mitoura)
especially where it is sympatric with utilized J. virginiana or J. scopu-

16 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

lorum. Johnson & Borgo (1976) have shown that the perching behavior
of C. siva and C. gryneus is distinctly patterned and preferenced for
heights. They postulate that the nature of this patterned perching be-
havior selects against prostrate morphs and is at least a partial boundary
on their usage as a larval foodplant (Johnson & Borgo, 1977). The im-
portance of the number of needled leaves on both J. horizontalis and J.
communis needs investigation since first instar larvae burrow into these
to feed.

Knowledge of the local specificities of the two oligophagous species
is quite incomplete, although preliminary evidence from several localities
indicates that populations are specific to particular plant species. In Palo
Duro Canyon (Randall and Armstrong cos., Texas) J. scopulorum, J.
pinchotii, J. monosperma, and hybrids of the latter two occur (Adams,
1972, and pers. comm.). Field data from collectors of C. gryneus indicate
that J. pinchotii is the only foodplant. However, verification is needed
by someone who can test this hypothesis directly. Peter Herlan (pers.
comm.) reports that the Great Basin brown morph of C. siva feeds ex-
clusively on J. osteosperma. Perhaps this is true, but Vasek (1966) has
suggested that this species introgresses with J. occidentalis westward,
and the taxonomic relationships of C. siva in the northwest basin are
now indicated as including two, largely disjunct subspecies, one feeding
on J. occidentalis in central and eastern Oregon and the other on J.
osteosperma in Nevada eastward to Utah. In Missouri and Arkansas,
C. gryneus populations are located on J. ashei where it occurs as “islands”
within the range of J. virginiana. Other C. gryneus populations are on
J. virginiana. This is another location ideal for specificity studies, as are
the areas of diversity of juniper species in Arizona and New Mexico. In
California, John Lane reports (pers. comm.) C. siva juniperaria perching
on both J. osteosperma and J. occidentalis in an area where J. occidentalis
has been reported as the foodplant. Thus, foodplant relations in C. siva
and C. gryneus mirror situations reported in Burns (1964), Downey
(1966), and Downey & Dunn (1965). Local specificities are due to
oviposition by the female on the plant species it fed on as a larva. Thus,
according to the familiar “Hopkins’ Host Principle,” specificity is main-
tained. However, it is obvious that alterations do occur through time
and space (as the above authors also indicate), and this is why such
species show catholicity when their foodplant usage is viewed as a whole.
The mechanism of ovipositional specificity and the nature of chance
alterations need further elucidation. Downey & Dunn (1965) suggest
that the patterning of Hopkins’ Host Principle is not genetic but physio-
logical and undergoes divergence, convergence, and parallelism through

VoLUME 32, NUMBER 1 17

time and space. The present study indicates that similar foodplants
available as replacers offer opportunity for divergence, since nearly all
barriers posed to these insects by replacer plants have been crossed.
Similarly, the remarkable coincidence of distinguishable morphs or sub-
species generally within the distribution of one or another foodplant or
foodplant relative suggests that foodplant adaptations play an important
role in subspeciation. Callophrys gryneus sweadneri inhabits the areas
of J. silicicola, C. gryneus gryneus those of J. virginiana, and C. gryneus
castalis those of the transition of the latter plant to the ranges of J. ashei
and J. pinchotii. Relations in the C. siva complex, although trinomial
knowledge is less complete, are equally distinctive. If one assumes
monophagous C. hesseli evolved through adaptations of some populations
of early C. gryneus stock to Chamaecyparis thyoides, a similar mechanism
is imaginable, especially since C. thyoides and J. virginiana have under-
gone a change in their degree of sympatry through time (M. Rosenzweig,
pers. comm.) in which populations of C. thyoides are now somewhat
disjunct, and those of C. hesseli apparently extremely so.

Laboratory foodplant experiments with these species have not been
extensive, and such data is of limited use in drawing inferences about
foodplant utilization or preference in nature (Downey & Dunn, 1965;
Downey & Fuller, 1962). However, studies to date indicate that quite
divergent Cupressaceae species are at least nutritionally adequate and
otherwise edible by some of the Callophrys (Mitoura) species. There is
a need to further clarify the reported acceptance of J. virginiana by
larvae of C. hesseli.

Distributional relations. One comment on the distribution of these
insects, with regard to the frequency of transplanted populations is ap-
propriate at this time. Cupressaceae species are widely used both in
agricultural and landscape planting, and a number of transplanted
Callophrys (Mitoura) populations have been noted (Figs. 13 & 14).
Taxonomists should be especially aware of this when studying the com-
parative morphology of these butterflies. The occurrence of C. siva in
planted forest well isolated in central Nebraska, where juniper is raised
from Rocky Mountain stock, is an extreme example, as is the occurrence
of this insect in a shelter belt along the Missouri River.

ACKNOWLEDGMENTS

Many lepidopterists and botanists contributed data for this study. I
owe a special debt to John Lane, Oakley Shields, and Arthur M. Shapiro
(Univ. of California, Davis) for discussing Callophrys (Mitoura) species
with me, and to Frederick H. Rindge (American Museum of Natural

18 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

History, New York) for providing the facilities of the museum and
editing this paper. For general support I am grateful to the staff of the
Museum of Natural History, University of Wisconsin, Stevens Point,
especially Charles A. Long, Director, and Robert Freckmann, Curator of
the Herbarium. Debt is owed John C. Downey (University of Northern
Iowa, Cedar Falls ) who introduced me to this group and suggested topics
for research. The following provided data for which I am very grateful:
R. A. Anderson, J. F. Gates Clarke, Ernst J. Dornfield, J. Donald Eff,
George Ehle, Scott Ellis, Frank D. Fee, Clifford D. Ferris, Mike Fisher,
Richard Funk, Richard Guppy, Lucien Harris, J. Richard Heitzman,
Peter Herlan, Sidney Hessel, Ronald Hooper, Roderick R. Irwin, Roy O.
Kendall, H. L. King, Alexander B. Klots, Steve Kohler, Henry A. LeBeau,
Bryant Mather, James R. Maudsley, Tim McCabe, Lee D. Miller, James
Mori, John S. Nordin, Roger Pease, Richard Priestaf, Kilian Roever,
Michael Rosenzweig, R. E. Sanford, James Scott, Ernest M. Shull, William
Sieker, Michael J. Smith, J. Bolling Sullivan III, Fred Thorne, and J. W.
Tilden.

I am indebted to the herbaria at Arizona State University (Tempe);
Colorado University (Boulder); University of Florida (Gainesville); Uni-
versity of Mexico (Mexico City); University of Saskatchewan (Regina);
and University of Wisconsin (Stevens Point) and especially to Cupres-
saceae specialists R. P. Adams (Colorado State Univ., Fort Collins) and
David F. Van Haverbeke (University of Nebraska, Lincoln) for their

comments and aid.

LITERATURE CITED

ApaMs, R. P. 1972. Chaemosystematic and numerical studies of natural popula-
tions of Juniperus pinchotii Sudw. Taxon 21: 407-427.

Apams, R. P. & B. L. Turner. 1970. Chaemosystematic and numerical studies
of natural populations of Juniperus ashei Buch. Taxon 19: 728-751.

ANDERSON, R. A. 1974. Southern records of Mitoura hesseli (Lycaenidae). J.
Lepid. Soc. 28: 161.

Brower, L. P. 1958. Larval foodplant specificity in butterflies of the Papilio
glaucus group. Lepid. News 12: 103-114.

Brown, F. M., D. Err, & B. Rorcer. 1957. Colorado butterflies. Denver Mus.
Natur. Hist., Denver, 368 p.

Burns, J. M. 1964. Evolution in skipper butterflies of the genus Erynnis. Univ.
Calif. Publ. Ent. 27: 1-216.

Comstock, J. A. 1927. Butterflies of California. By author. 344 p.

Downey, J. C. 1966. Host-plant relations as data for butterfly classification.
System. Zool. 11: 150-159.

Downey, J. C. & D. B. Dunn. 1965. Variation in the lycaenid butterfly Plebejus
icarioides III. Additional data on foodplant specificity. Ecology 45: 172-178.

Downey, J. C. & W. C. Futter. 1962. Variation in Plebejus icarioides (Lycae-
nidae ). I. Food-plant specificity. J. Lepid. Soc. 15: 34-42.

EMMEL, T. C. & J. F. Emmet. 1973. The butterflies of southern California. Nat.
Hist. Mus. Los Angeles Co. Sci. Ser. 26: 1-148 p.

VOLUME 32, NUMBER 1 19

Fassett, N. C. 1945. Juniperus virginiana, J. horizontalis, and J. scopulorum—
IV. Hybrid swarms of J. virginiana and J. horizontalis. Bull. Tor. Bot. Club
72: 379-384.

Jounson, K. 1972. Juniperus (Cupressaceae) speciation and the ranges and
evolution of two Callophrys (Lycaenidae). J. Lepid. Soc. 26: 112-116.

Jounson, K. 1976a. Three new Nearctic species of Callophrys (Mitoura), with
a diagnostis (sic) of all Nearctic consubgeners (Lepidoptera: Lycaenidae).
Bull. Allyn. Mus. No. 38, 30 p.

1976b. A new species of Callophrys (Mitoura) from Mexico (Lepidop-

tera: Lycaenidae). Pan-Pacific Ent. 52: 60-62.

Callophrys (Mitoura) nelsoni (Boisduval) and siva (Edwards) in the
northwestern United States, Lycaenidae. (Ms. in prep.).

Jounson, K. & P. M. Borco. 1976. Patterned perching behavior in two Callophrys
(Mitoura), Lycaenidae. J. Lepid. Soc. 30: 169-183.

Perching behavior as a partial boundary of distribution and foodplant
utilization in Callophrys (Mitoura) (Lepidoptera: Lycaenidae). (in review).
Kxuors, A. B. 1951. A field guide to the butterflies of North America, east of the

Great Plains. Houghton Mifflin Co., Boston, xvi + 349 p.

LirtLe, J. B. 1971. Atlas of United States trees. U.S.D.A. Misc. Publ. No. 1146.

Pease, R. W. 1963. Extension of known range of Mitoura hesseli. J. Lepid.
Shen a Romer a

Rawson, G. W., J. B. Zrecier, & S. A. Hesset. 1951. The immature states of
Mitoura hesseli Rawson & Ziegler. Bull. Brooklyn Ent. Soc. 46: 123-130.

Remincton, C. L. & R. W. Pease. 1955. Studies in foodplant specificity. 1.
The suitability of Swamp White Cedar for Mitoura gryneus (Lycaenidae).
Lepid. News. 9: 4-6.

Ross, J. G. & R. E. Duncan. 1949. Cytological evidences of hybridization be-
tween Juniperus virginiana and J. horizontalis. Bull. Tor. Bot. Club 76: 419-
429. -

ScHurtz, D. L. 1971. A tri-parental swarm of Juniperus L. Unpubl. PhD Thesis,
University of Nebraska, Lincoln. 234 p.

U.S.D.A. 1953. Checklist of native and naturalized trees of the United States
(including Alaska). U.S.D.A. Agric. Handb. No. 41. 472 p.

Van HaversBEKE, D. F. 1968. A population analysis of Juniperus in the Missouri
River Basin. Univ. Nebr. Studies, No. 38. 82 p.

Vasex, F. C. 1966. Distribution and taxonomy of three western Juniperus. Brit-
tonia 18: 350-372.

Journal of the Lepidopterists’ Society
32(1), 1978, 19

IMPORTANT NOTICE TO CONTRIBUTORS

Authors should submit an abstract for all articles to be published in the Journal
beginning with Volume 32, issue No. 1. The abstract should summarize the im-
portant contents and conclusions of the paper in concise and specific sentences. It
should not exceed 1-3% of the paper's total length, and should indicate the ob-
jectives, methods, and topics covered by the paper. References to literature, illustra-
tions, and tables should be omitted from the abstract, since this should be com-
pletely self-explanatory. These abstracts will be printed at the front of each paper,
and also will be forwarded to Biological Abstracts for possible inclusion therein.
Such abstracts should accompany articles only. They are not required for general notes.
Authors who now have papers in press should forward such abstracts to the new
editor at their earliest convenience.

Journal of the Lepidopterists’ Society
32(1), 1978, 20-36

FOODPLANT, HABITAT, AND RANGE OF
CELASTRINA EBENINA (LYCAENIDAE)

WarRREN HERB WAGNER, JR.
Department of Botany, The University of Michigan, Ann Arbor 48109

T. LAwRENCE MELLICHAMP
Department of Biology, University of North Carolina, Charlotte 28233

ABSTRACT. The larval foodplant of the recently described Celastrina ebenina
Clench is Aruncus dioicus (Walt.) Fernald (Rosaceae), the Goat’s-beard. Over
150 adults were raised from eggs and young larvae. The range of the butterfly
coincides nicely with that of the plant, from Pennsylvania to North Carolina and
Missouri. The habitat for plant and butterfly is moist, rich forest. The closely
related C. pseudargiolus Boisduval & LeConte is vastly more abundant and ubiq-
uitous than W. ebenina, and has a wide variety of larval foodplants. Larvae of
C. ebenina differ in several respects from those of C. pseudargiolus, including color
pattern and stellate processes. Also described and discussed are the plant and
butterfly associates of C. ebenina, flower visitations of the adults, experiments on
foodplant specificity, feeding characteristics of the larvae, broods, botany of the
foodplant, and geographical distributions, including a number of new locality
records. A guide for discovering new colonies of this rare eastern American butter-
fly is provided.

Except for brief reports (Clench, 1972; Wagner & Showalter, 1976),
little has been published on the biology of the poorly known Dusky Blue
Butterfly, Celastrina ebenina Clench, of the eastern United States. This
lycaenid is notable for several reasons. Interpreted for over a century
as an aberration or form, it was not recognized as a distinct species until
1972. The colors of the upper surfaces of the males and females are
peculiar for being the reverse of the usual situation among plebejine
blues in that the males are dull, dark grayish-brown or blackish, while
the females are mainly lustrous blue. The insect is regarded as especially
rare and local, having been reported previously, usually as just one or
a few individuals, from only 12 localities. Knowledge of its foodplant,
behavior, habitat, and geographical distribution has been incomplete or
lacking.

The present paper records the results of research in 1976. We now
understand the ecology of C. ebenina far better than we did in the past,
and we believe that we have an explanation for the geographical distri-
bution and sporadic occurrence of the species. At the outset of this study,
as botanists, we entertained the possibility that the peculiarities of oc-
currence of C, ebenina might be due to specialized larval foodplant
preference,

Because of earlier reports of the species there, the area chosen for
our field investigations was in the Daniel Boone National Forest, in and

VoLUME 32, NUMBER 1 2]

around the Red River Gorge in Powell and Menifee cos., Kentucky, a
few miles north of the town of Slade. We found that habitats suitable
for C. ebenina occur extensively, though sporadically, throughout this
area, and in some places the butterfly is common or abundant, though
extremely localized, flying with the much more numerous and ubiquitous
Common Blue or “Spring Azure,” C. pseudargiolus Boisduval & LeConte.
The topography in the Red River Gorge is made up of steep, abrupt hills
and valleys, ranging from 700’ altitude in the river and stream beds to
over 1300’ at the tops of the highest hills. Commanding cliffs of light
tan or whitish sandstone crop out at the crests of some of the hills, but
in the valleys where C. ebenina flies, the most conspicuous rock is a
loose, broken, dark-gray shale. The general area is mainly traversed by
narrow dirt roads, especially along the larger streams, and there are
only occasional two-lane hardtop roads.

Our search for suitable habitats to study was initiated on 17 April 1976.
We drove from place to place along the country roads, stopping wherever
we encountered roadside puddles or wet streamside flats. In such spots
accumulations of butterflies were the rule when it was bright and sunny,
especially between the hours of 0900 and 1400. Among the guests at
these “puddle parties” we encountered rare males of C. ebenina, prac-
tically always with at least several and usually many C. pseudargiolus,
the latter being much more conspicuous in flight. In spite of our success
in encountering specimens of C. ebenina here and there over an area of
perhaps a dozen square miles, at no place were there more than a few
individuals. Rare observations of females showed them almost always to
be in flight, and following them gave no clue to where their eggs were
laid.

Finally, at around 1030 hrs the following day, we came upon an excel-
lent locality—a moist, steep, rocky, north-facing wooded slope, along a
narrow dirt road along the Red River. In only 20 min, ca. three dozen C.
ebenina of both sexes were observed, the females numbering about twice
as many as the males. The latter were all more or less worn, dull in
appearance, and flying along the edges of the road, occasionally alighting
on wet muddy spots (Fig. 1, lower photo). Most of the females,
however, were in fresh condition and were flitting around the vegetation
on the slopes above and below the road. It was obvious that many of
them were engaged in oviposition. They flew rapidly in an “exploratory”
pattern, pausing often at a single species of plant, the Goat’s-beard,
Aruncus dioicus (Walt.) Fernald. Occasionally the females landed on
the abundant and conspicuous Wild Hydrangea, Hydrangea arborescens
L., but usually only momentarily.

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

bo
bo

Fig. 1. Celastrina ebenina habitat along Red River, Powell Co., Ky. Upper:
View upstream showing extensive understory growth on forest slope. Butterflies
visit Geranium flowers here. Lower: View downstream showing damp ruts in road
where males congregate. (Photo by J. M. Beitel. )

VOLUME 32, NUMBER 1 23

Figs. 2-6. Celastrina ebenina adults and eggs: 2, female laying eggs on young
Aruncus shoots (R. P. Carr); 3, freshly emerged male, showing blue scaling (T. L.
Mellichamp):; 4, eggs on leaf and inflorescence primordia of Aruncus (cf. fig. 14)
(T. L. Mellichamp): 5, unhatched eggs; 6, eggs (two of them hatched) showing
wall pattern detail (R. P. Carr).

On Aruncus the butterflies alighted on very young, unfolding leaflets,
and then walked around slowly, laying eggs (Fig. 2). After an individ-
ual would fly away, we could easily find the eggs, mainly on the lower
blade surfaces, on and between the main veins of the leaflets (Fig. 4).
At this time of year the main axis of the plant is still embryonic, and the
habit is very different from the mature habit with the inflorescence fully
developed (cf. Fig. 13, full-grown plant, and Fig. 14, stage at time of
oviposition). When freshly laid, the eggs showed a grayish blue color.

24 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Sometimes several eggs are laid in the same spot, but usually they are
laid separately (Figs. 4, 5, & 6). Egg-laying occurred over 1% hours of
observation, and there was no sign of abatement after 1200 hrs when we
left the site.

For careful observations, a total of 18 young cuttings like those in Fig.
14 were randomly collected. The two oldest leaves overtop the main
shoot at this stage. The main shoot, with its very young leaves and
inflorescence primordium, is only about one-fourth the length of the
oldest leaf (the large, bipinnate leaf on the right side of the figure) and
one-half the length of the next oldest leaf. The softest, most embryonic
tissues are those of the primordial main shoot. We found a total of 133
eggs altogether on the collected shoots—35 on the oldest leaves, 26 on
the next oldest, and 66 on the young main shoots. Thus, our evidence
suggests that the butterflies prefer to lay eggs on the youngest tissues.
The number of eggs averaged seven per cutting, but one had 18.

Naturally we wondered whether at least some of the eggs we found
did not represent the closely related C. pseudargiolus, which flies in
large numbers at this locality, a species noted for its polyphagy. There-
fore, we decided a couple of weeks later to conduct an experiment de-
signed to help us answer this question, as will be described below.

The plant community on this slope is a rich mixed-mesophytic forest.
We recorded a total of 31 trees and shrubs and 56 herbs (including ferns
and graminoids ) in the area where C. ebenina was ovipositing. Most of
these plants are typical associates of Aruncus dioicus, and some of the
more prominent ones will be enumerated later in our discussion of this
plant.

The best place to find C. ebenina adults is in association with other
mud-loving butterflies (C. pseudargiolus, Callophrys henrici, Erynnis
spp., and Papilio spp.) in damp spots along dirt roads and gravelly,
sandy, or muddy river flats. Practically all of the “mudding” individuals
we observed were males, sometimes as much as a quarter of a mile from
the foodplant, although usually much closer. On only two occasions did
we find females landing on wet soil. We disturbed one of them several
times, but each time it returned.

The only flower which seemed to attract C. ebenina at this locality
was the Wild Geranium, Geranium maculatum. The showy rose-purple
flower has a flat five-petalled corolla 3.0-3.5 cm across. Bearded nectaries
occur between and at the bases of the petals. The butterflies walk over
the top surfaces of the corolla and probe between the petal bases. Later

'A complete list of the associated plants at this locality will be sent upon request to readers.

VOLUME 32, NUMBER 1 25

Figs. 7-12. Celastrina ebenina immature stages: 7 & 8, instar 1 caterpillars (J.
G. Bruce III); 9 & 10, instar 2 caterpillars (J. G. Bruce III); 11, mature caterpillars,
showing pale, poorly contrasting pattern and characteristic leaf damage; 12, pupa
attached to Aruncus leaf (R. P. Carr).

we discovered that the bulk of individuals obtain their nectar from
underneath the flower! Both sexes flit from flower to flower, landing
on the peduncle or on the underside of the perianth, then walking toward
the sepal bases where they insert their proboscises. So positioned on the
flowers, the butterflies are invisible from above.”
We recorded all of the species of butterflies we found in association
* Curious to see whether other butterflies behaved in the same manner, we discovered (in

Michigan in the middle of May) that Erynnis juvenalis displayed the same routine on Geranium
flowers.

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

pecaaimnnanat e —.rrtrs——C

1O 6M.

(‘a

—s

13, habit drawing of fully grown Aruncus dioicus showing inflorescence (apex) and form
y); 14, young shoot of Aruncus at time of oviposition by C. ebenina (see text) (T. L.

~, Vg ,
Vial of
WEN Nia
Fog a
; AS .
RSS
Cor )
A neil

Figs. 13-14. Celastrina ebenina foodplant:
of compound leaves in mid-June (Del. J. G. Lac

Mellichamp ).

VoLUME 32, NuMBER 1 27

ee

= *\\

Fig. 15. Geographical distribution of Aruncus dioicus (stippling) and docu-
mented localities for Celastrina ebenina (dots) (T. L. Mellichamp).

28 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

with C. ebenina on 18 April. The following list is a compilation, since no
one locality had all of the species (those marked with an asterisk oc-
curred with C. ebenina on wet soil): Papilio glaucus,* P. troilus,*
Battus philenor, Pieris virginiensis,* Colias eurytheme, C. philodice,
Polygonia comma, P. interrogationis, Boloria bellona, Phyciodes tharos,*
Callophrys henrici,* Celastrina pseudargiolus,* Erynnis brizo,* E.
juvenalis,* Amblyscirtes samoset, A. aesculapius, and Epargyreus clarus.
On several later visits to the study area by Loran D. Gibson, Amos H.
Showalter, and ourselves, the following additional species were observed,
most of them, however, well beyond the normal flight period of C.
ebenina, which ended the first week of May: Limenitis astyanax,
Speyeria cybele, Asterocampa celtis, Ewptychia hermes, E. cymela, Lethe ~
creola, L. anthedon, Autochton cellus, Thorybes pylades, and Poanes
hobomok.

Experiment on Foodplant Specificity

The question we addressed ourselves to was whether C. pseudargiolus
also shared the foodplant of C. ebenina. The former is abundant in the
Red River Gorge area and is in constant association everywhere with C.
ebenina. Although we did not observe C. pseudargiolus ovipositing upon
Aruncus, this would not preclude the possibility that it occurs. The num-
ber of Celastrina eggs we observed was so great that it seemed reasonable
to assume, because of their morphological similarity, that perhaps some
of them belonged to the abundant species, especially since C. pseu-
dargiolus is noted for its varied bill of fare. Literature records show that
no less than 10 families of flowering plants contain larval foodplants for
this species, and we have discovered early stages of it on such different
families as Caprifoliaceae and Cornaceae. It would not have surprised
us, therefore, to find C. pseudargiolus utilizing Aruncus, especially since
its larvae have been reported on Spiraea, another closely related genus
in the same subfamily of Rosaceae.

On 2 May, accordingly, we revisited the Red River Gorge area and
collected from seven to nine shoots with attached eggs and minute larvae
from each of five colonies of the foodplant, all of them a mile or so
separate from one another. These shoots were brought back to The Uni-
versity of Michigan in Ann Arbor, and the eggs and young larvae were
raised to adulthood. Extra foodplant shoots were kept in plastic bags
and refrigerated to preserve them until they were needed, and additional
foodplants were grown for further supplies. The cut bases of the shoots
of Aruncus bearing eggs and larvae were inserted in jars of water in
| x 1 x 2 ft glass aquaria covered with a plastic material to keep the

VoLUME 32, NUMBER 1 29

larvae from escaping. Some of the caterpillars drowned when they
walked down the main stems into the water, others escaped and were
lost, and some died as a result of cannibalism. Nevertheless, a total of
153 butterflies were raised to maturity by the end of the second week
in June, 76 males and 77 females. Another eight butterflies came from
larvae which escaped from their containers. The first caterpillars
pupated on 16 May, 14 days after the field collection, and the first adults
to emerge appeared on 23 May. This second brood apparently does not
occur in nature (see below).

The results of this experiment were striking. Not only were none of
the “checkery” caterpillars of C. pseudargiolus noticed among the paler,
less contrasty larvae of C. ebenina (Figs. 7-9), but not a single one of
the adults was C. pseudargiolus; all were C. ebenina. Furthermore, the
14 additional butterflies from our 18 April eggs were all C. ebenina.
Also, the four individuals that emerged from dormant chrysalids the
following winter were all C. ebenina. With C. pseudargiolus as abundant
as it is in the localities where our collections were made, one would
expect at least some evidence of its occurrence on Aruncus if it fed on
that plant at all.

It is interesting to note that ca. one-third of the males display blue
scales (Fig. 3), at least at the time of emergence. Although the original
description by Clench (1972, p. 37) records the male upperside as
“uniform blackish brown when fresh,’ blue scales are conspicuous in
these individuals, especially on veins R, to R; and the discal portions of
M;, Cu,, and Cu, on the fore wings, and scattered in the discal area of
the hind wings. The occurrence of blue scaling on the males may con-
stitute a regional difference. The ground color above of the males varies
from blackish gray to pale slate gray. The females vary greatly in ground
color (whitish blue to fairly intense blue) and the amount of dark mark-
ing. There is no confusing any of these specimens, however, with any of
the forms of C. pseudargiolus known to us.

To make preliminary comparisons of the caterpillars of C. ebenina
and C. pseudargiolus, we obtained eggs and first instar larvae of the
latter in two localities near Ann Arbor in Washtenaw Co., Michigan.
They were readily found on the young inflorescences of Grey Dogwood,
Cornus racemosa and Red Osier, C. stolonifera (Cornaceae), and Nanny-
berry, Viburnum lentago (Caprifoliaceae); the dates of collection were
8 and 22 May. We maintained the larvae under the same conditions as
those of C. ebenina but kept them in a separate room to avoid any op-
portunity of possible escapees getting mixed up.

We did not make detailed comparisons of the larval morphology, but

30 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

certain differences were obvious. The final instar caterpillars of C.
ebenina (Fig. 11) are considerably more uniform in color than those of
C. pseudargiolus. They are pale whitish blue-green, with three slightly
contrasting longitudinal stripes of yellowish white, two lateral and one
dorsal. Between the dorsal and each lateral stripe is a line of yellow—
white dots, and similar dots and dashes are scattered over the body. The
caterpillars of C. pseudargiolus, as is well known, are more variable in
eround color, ranging from pinkish or pale bronze to yellowish green;
the pattern is much more conspicuously blotched as a rule. Earlier larval
stages of the two species (young instars of C. ebenina shown in Figs.
7-10) are very similar in their uniform pale green color.

In both species the mature larvae are velvety because of elaborate
stellate processes, each with a narrow and pointed filament arising from
the center. In C. ebenina the filaments are only slightly curved and
stand nearly erect. In C. pseudargiolus the filaments are more strongly
curved and tend to be arched over such that they are distally nearly
parallel to the body surface. This difference applies mainly to the more
abundant, smaller filaments; larger ones in both species are more alike,
only slightly curved and nearly erect.

Another difference between the larvae of the Dusky and Common
Blues involves the use of their respective foodplants. Although larvae of
C. ebenina occasionaily feed upon the tiny embryonic floral primordia
of Aruncus, the bulk of their feeding is upon the blade tissue between
the major lateral veins of the leaflets. Their feeding produces charac-
teristic elongated perforations in the blades; these persist when the
leaflets have achieved their full size, and give valuable clues during late
spring and summer for localities of the butterfly. Fig. 11 shows the
characteristic perforations. The younger caterpillars were observed on
the tops of the leaves more often than the older ones. Feeding between
a pair of lateral veins usually begins near the midrib of the leaflet and
progresses outward, but only rarely all the way to the margin and in-
cluding it. Very large caterpillars in the last instar may eat major lateral
veins.

In our experience the larvae of C. pseudargiolus feed primarily upon
floral primordia. Their eggs and larvae are found in inflorescenses. We
tried moving larvae to leaves, but always, after eating a small amount,
they would return to their floral clusters where they continued feeding.

Larvae of C. pseudargiolus bite into the sides of the closed flower bud
or the inferior ovary and eat the entire contents or leave certain parts

(e.g., petals). The body of the caterpillar sits motionless on the floral
pedicel or the side of the bud, and the extensible head is projected into

VOLUME 32, NuMBER 1 31

the cavity in the young flower to feed. From certain angles the larva
appears headless.

Broods

Previous field experience of our own and others suggested either that
C. ebenina is univoltine (in the Red River Gorge, flying from the second
week of April to the first week of May), or else, if it has more than one
brood, that adults of any later broods are so similar to those of C. pseu-
dargiolus that they have not been recognized as distinct. The second lab-
oratory brood is like the first brood. As it turned out, the vast majority of
our pupae (Fig. 12) emerged within a week or two of pupation. Of
those that failed to emerge and were kept until the following winter,
only a few produced butterflies.

The first adult from our 18 April field collections of C. ebenina eggs
emerged on 11 May. The first from our 2 May collections of eggs and
young larvae emerged on 23 May. Emergence continued until 29 May,
when it was stopped by placing the cultures in a coldroom from this
date until 4 June to keep more butterflies from appearing while we were
out of town. After we returned them to normal temperatures, emergence
resumed on 5 June and continued to 12 June. A grand total of 175 butter-
flies in our laboratory cultures seemed to demonstrate that there is a
second brood in the wild that follows closely upon the first, the second
brood flying from the second week in May to the first or second week
in June.

We therefore returned to the Red River Gorge to determine whether
a second brood occurred in nature. The results of our survey were most
unexpected. Loran D. Gibson visited there on 26 May and saw not a
single C. ebenina, although many other butterfly species were seen (in
litt., 2 June 1976). We then reconnoitered the area on 4 June, when the
butterflies should have been at their peak abundance, if there is a second
brood in nature. We saw no C. ebenina despite the fact that C. pseu-
dargiolus was common as well as 16 other species of butterflies. On 9
June, Amos H. Showalter searched the area and reported that “C. pseu-
dargiolus was common, but no ebenina” (in litt., 28 June).

After 13 June, the 28 pupae that remained in our cultures apparently
went into dormancy, and no more butterflies emerged. At the end of
June, therefore, we placed them in a coldroom at a temperature of 2°C
and left them there until 22 December. We hope that by keeping them
thus, in a set-up that we have used for “winterizing” fruits and seeds,
we might avoid the destructive effects other workers have had with C.
pseudargiolus, which involve either drying out or molding of the chrys-

32 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

alids. On 1 January 1977, two females emerged, one of which failed to
expand its wings. Another female emerged on 4 January, and still an-
other female failed to escape from the pupal skin. All the remaining
pupae appeared to have died, eight of them having moldy surfaces and
the rest having an unnatural brown color. }

What can we conclude regarding the broods of C. ebenina? In the
field we found no evidence for a second brood. If there is one, it must
be in extremely low numbers, i.e., a small “partial” brood. Somehow the
conditions of our laboratory cultures must have caused an abnormal
eclosion without the customary prolonged dormancy period. Some dia-
pause stimulus that effects C. ebenina must have been weak or missing
under the conditions of our experiment, and thus only a small percentage
of the chrysalids went into long-term dormancy.

Botany of the Foodplant

The colloquial name of the foodplant, Aruncus dioicus, may cause
some confusion, since its name “Goat’s-beard” is applied also to the un-
related Tragopogon pratensis L. in the Asteraceae, a naturalized weed
from Europe. The generic name Aruncus comes from the Greek and
means literally “goat’s beard.” It is a member of the Rosaceae and is a
native eastern American plant of rich, mature forests. It is famous among
United States’ botanists as an illustration of convergent evolution, because
superticially A. dioicus resembles closely the “False Goat’s-beard,” Astilbe
biternata, of the Saxifragaceae. So closely do these plants resemble each
other that they are regularly confused, even in herbaria. Ecologically
the two look-alikes occupy almost identical niches, and they are both
unusual among members of the mesophytic forest association in being
dioecious (male and female flowers being borne upon separate plants).
They are pollinated not by wind, which is the usual situation in dioecious
plants, but rather by insects, mainly small Hymenoptera. The geograph-
ical range of Aruncus dioicus is shown in Fig. 10. The range of Astilbe
biternata is much narrower, mainly in the mountains of North Carolina
and adjacent parts of Virginia, W. Virginia, Kentucky, Tennessee, South
Carolina, and Georgia. Aruncus overlaps it completely, so that students
of C. ebenina must be warmed of the danger of confusing the “True”
with the “False” Goat’s-beards in the area of their sympatry. Accord-
ingly, we have prepared a comparison of the two in Table 1, the most
obvious characters marked with asterisks. A line drawing of a mature
specimen of Aruncus nearly 1’4 m tall is reproduced in Fig. 1. The stage
of growth when the plant serves as larval food for C. ebenina is shown
in Fig. 2, corresponding to only the two bottom leaves and the lower

VOLUME 32, NuMBER 1] Oo

TaBLE 1. Comparison of “True” and “False” Goat’s-beards.

Aruncus dioicus Astilbe biternatum
*1. Stipules Absent Present
2. Terminal leaflet Unlobed 3-lobed
3. Leaf base Attenuate Cordate (heart-shaped )
*4. Veins per leaflet 8-18 pairs 8 or less pairs
*5. Leaf and stem hairs Absent Abundant ( glandular )
6. Marginal teeth Convex Acuminate
7. Sepals per flower 5 5)
8. Petals per flower 5B 0-5
9. Stamens per flower 15-20 8-10
*10. Carpels per flower 3-4 2

sixth of the drawing. The tissues upon which the caterpillars feed are
soft, and the earliest instars feed upon the most embryonic parts. The
plants grow rapidly and come into flower 5-7 weeks after the butterflies
lays their eggs.

Both Aruncus and Astilbe have rather massive underground stems that
produce large roots or root masses which hold the plants firmly in place
on steep slopes (Fig. 3). Many buds are present at the ground level,
and some of these may develop into shoots at the crown, producing
clumps of as many as eight flowering shoots. Their spreading compound
leaves fill in the space where they grow, presumably allowing little
growth of other plants beneath them.

The Goat’s-beards are most typical of rich, mesic woods, partly shaded
roadsides, and sloping sides of streams and rivers. The ancient habitat
was probably on steep eroding slopes and stream banks in dark forested
areas, but man has stimulated its spread by creating new habitats where
roads have been cut through the mesic forest. In a uniformly shaded
forest stand only 10% of the plants may flower, but when released from
the effects of low light levels, as on road cuts at the forest edge or steep,
eroding stream banks, the populations may display up to 100% flowering.
The plants require, however, relatively cool, moist conditions, and they
exist almost exclusively upon north-facing slopes. Aruncus is known to
occur as high as 5500’ altitude in the mountains (Buncombe Co., N.C.),
although the average occurrence throughout its range is considerably
less than 2500”.

Would-be collectors of C. ebenina should seek the foregoing site con-
ditions, with the following array of associated species (based upon
studies of a number of Aruncus localities by Mellichamp 1976): woody
plants—Acer rubrum, A. saccharum, Aesculus octandra, Betula lenta,
Carpinus caroliniana, Cornus florida, Fagus grandifolia, Lindera benzoin,

34 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Liriodendron tulipifera, and Tilia americana; herbs—Adiantum pedatum,
Athyrium filix-femina, Botrychium virginianum, Carex plantaginea,
Cimicifuga racemosa, Geranium maculatum, Impatiens capensis, La-
portea canadensis, Tiarella cordifolia, and Trillium spp. The presence
of a majority of these species, together with Wild Hydrangea, Hydrangea
arborescens, given the topographical conditions cited, especially a north-
facing slope, should lead to colonies of Aruncus and therefore C. ebenina.

Ranges of Plant and Butterfly

There is a remarkable correlation between the known localities where
the butterfly has been found and the geographical distribution of Aruncus
dioicus. Clench (1972) has already discussed doubtful records for C.
ebenina, including New York City and southern Colorado. We should
like to add to the list of doubtful records that of Blatchley in Wabash
Co., Indiana (Clench, 1972, p. 41), unless an actual specimen is dis-
covered. The “black male” he referred to could have been a melanistic
male of C. pseudargiolus, a wind-blown stray of C. ebenina, or a mis-
labeled specimen. Clench mentions that Edwards had vaguely attributed
the species to Tennessee and Georgia, although Clench himself had seen
no specimens in these areas. It now seems very likely, as Clench (1972)
suggested, that C. ebenina may turn up in both of those states.

Aruncus grows far to the west of the localities cited in earlier studies
of C. ebenina (Clench, 1972; Wagner & Showalter, 1976), the western-
most documented records for which were all east of Cincinnati, Ohio, and
Lexington, Kentucky. We can now report records in Illinois, Missouri,
and Arkansas. Charles L. Remington took a fresh male south of Elsah,
Jersey Co., Illinois, on 15 April 1942 (in litt., 21 Jan. 1977). He found
additional males in St. Louis Co., Missouri, in the late 1930’s. Also, J.
Richard Heitzman (in litt., 14 July 1976) has informed us that he ob-
tained by exchange “a male and a female taken 4 May 1924 at Creve
Coeur Lake near St. Louis, St. Louis Co. There is no collector’s name,
but it should have been one of the active collectors of the day, probably
Ernst Schwarz, E. P. Meiners, or H. I. O’Burme. AII were active at the
time and collected often at Creve Coeur.” The dot shown in Fig. 10 for
Arkansas is based upon a single male taken in Hickory Flat Hollow,
Washington Co., on 2 April 1973 by Edward Gage. Gage writes (in litt.,
Dec. 1976): “This male particularly stood out as it was flying about a
mud puddle with several C. argiolus and Everes comyntas. .. . I im-
mediately assumed that it was a melanic form and quickly collected it.
The site was in close proximity to a draw or shallow canyon. About
90% of the immediate surrounding area is deciduous hardwood. ... Can-

VoLUME 32, NUMBER 1 By5)

yons and woodland extend all around Beaver Lake from the collecting
site. ”

CONCLUSIONS

Celastrina ebenina may have larval foodplants other than Aruncus
dioicus. However, it should be noted that none of the unquestioned
localities of this butterfly lies outside the known range of Aruncus.
Furthermore, we now have good reason to believe that the abundant
Common Blue, C. pseudargiolus, does not share the foodplant of the
Dusky Blue, C. ebenina. Our experimental raising of eggs and young
larvae on Aruncus from areas in which C. pseudargiolus is abundant
revealed not a single specimen of that species. All were C. ebenina.

One reason for apparent rarity of C. ebenina in comparison with its
near relative is that its geographical range is much more limited. Another
is that it is probably monophagous rather than polyphagous. Its food-
plant is confined to one habitat—north-facing, richly wooded, shaded
slopes, so that the butterfly tends to be highly localized and colonial. The
multiple foodplants of C. pseudargiolus occupy many habitats, and the
butterfly is therefore practically ubiquitous.

Celastrina ebenina is probably often overlooked. The dull males in
flight may suggest badly worn individuals of C. pseudargiolus. Celastrina
ebenina tends to fly closer to the forest floor (a concomitant of its under-
story foodplant?), whereas C. pseudargiolus has a slow, up-and-down
flight reaching the shrub and lower tree layer. The bright reflecting blue
of C. pseudargiolus males plus their tremulous flight pattern through
the woods at heights of roughly 2-10’ make them especially visible. The
females of C. ebenina resemble dull females of C. pseudargiolus, but
they are even more localized than the males, rarely even visiting mud
puddles, occurring rather in the herb layer of the forest in more or less
checkered sunlight. We wonder how many collectors (including our-
selves!) in the spring have overlooked C. ebenina while they focused
instead on such critical genera as Erynnis and Callophrys as well as such
widely advertised rarities as Pieris virginiensis and Erora laeta, both of
which are now known to fly in association with C. ebenina.

If the conclusions of this research are correct, we predict that C.
ebenina will be found not only in many new localities in the states from
which it is already known, but, in addition, southern Indiana, eastern
Tennessee, northern Georgia, western South Carolina, and western Mary-
lan. To help achieve this, we propose the following formula:

1. Locate areas of rich mesophytic forest in rolling or mountainous
country.

36 JoURNAL OF THE LEPIDOPTERISTS SOCIETY

2. Follow roads or streams and find north-facing, cool, shaded forest
slopes with some erosion or disturbance.

3. Look for plant associations including such trees and shrubs as
Hydrangea, Acer, Aesculus, Betula, Carpinus, Cornus, Fagus, Lindera,
Liriodendron, Rhododendron, and Tilia plus the majority of herbs given
above.

4, Explore for large colonies of “Goat’s-beard,” Aruncus dioicus, the
larval foodplant, especially on north-facing roadsides and streamside
slopes.

5. Visit the area in April and early May in search of C. ebenina—
males on muddy spots, females around the foodplant, and both sexes on
Geranium flowers (careful!—they may be underneath the petals).

6. Or, if the weather is cloudy or rainy, search the young shoots of
Aruncus for greenish blue, rough-surfaced eggs, and (or) pale green
caterpillars, the latter evidenced by narrow perforations in the soft leaf
tissue between the veins of young leaflets.

To sum up, our evidence thus far indicates that Celastrina ebenina is
a “specialist,” not a “generalist.” When compared with C. pseudargiolus,
it has a narrow range (not broad), one foodplant (not many), a single
brood (not several), and an essentially uniform morphology (not many
varieties and forms ).

ACKNOWLEDGMENTS

We thank the following persons for their help in this project: J. M.
Beitel, J. G. Bruce III, Robert P. Carr, D. J. Harvey, Loran D. Gibson,
J. Richard Heitzman, Janice Glimn Lacy, Amos H. Showalter, Florence
S. Wagner, and K. S. Walter. We are especially grateful to Harry K.
Clench for his encouragement and critical advice.

LITERATURE CITED

CLeENCH, Harry K. 1972. Celastrina ebenina, a new species of Lycaenidae
(Lepidoptera) from the eastern United States. Ann. Carnegie Mus. 44: 33-44.

Me.iicuamp, T. L. 1976. A comparative study of Aruncus (Rosaceae) and
Astilbe (Saxifragaceae ), and the problem of their relationships. Doctoral Thesis.
The University of Michigan, Ann Arbor.

Wacner, W. H., Jr. & AMos H. SHowatrer. 1976 Ecological notes on Celastrina
ebenina (Lycaenidae). J. Lepid. Soc. 30: 310-312.

Journal of the Lepidopterists’ Society
32(1), 1978, 37-48

STUDIES ON RESTINGA BUTTERFLIES. II. NOTES ON THE
POPULATION STRUCTURE OF MENANDER FELSINA
(RIODINIDAE)

Curtis J. CALLAGHAN
IBM do Brasil Ltda., P.O. Box 1830, Rio de Janeirv, Brazil

ABSTRACT. The objective of the study was to describe various aspects of the
adult behaviour and population dynamics of the riodinid butterfly Menander felsina
(Hew). The conclusions were based on four years of field observations and a
marking-recapture study conducted over a period of 15 weeks.

The population was characterized by low intensive and extensive frequencies
(rarity in numbers and space respectively), characteristics shared by many other
forest riodinid species. M. felsinad maintained constant population levels of about
19 individuals over the 15 week marking-recapture period, due to 1) longevity above
that of most holarctic lycaenids, 2) low egg laying frequency both in time and space,
and 3) male territoriality, which results in older males doing most of the mating.
Limited adult and larval food sources were discounted as an explanation.

The population was found to be distributed in small groups or colonies near food
plant localities. The reasons for this were the low extensive distribution of foodplants
coupled with high female vagility. Depending upon conditions at each foodplant
locality, such as predation and exposure to the elements, each colony could become
extinct, only to be reestablished by another wandering female.

Studies on the ecology of neotropical butterflies to date have been
concerned for the most part with the larger species such as heliconiines
(Crane, 1955, 1957; Turner, 1971; Ehrlich & Gilbert, 1973), nymphalines
(Benson & Emmel, 1973; Young, 1972), and papilionids (Cook e¢ al.,
1971). Smaller butterflies such as riodinids have received only super-
ficial treatment, mainly consisting of scattered notes on observed habits
and population density. Early writers such as Bates (1864) and Seitz
(1913) made reference to the rarity of individuals and the great num-
bers of species that characterize neotropical riodinids. This has been
confirmed statistically by Ebert (1969), who used the terms low ex-
tensive and intensive frequency to denote the rarity of populations and
individuals, respectively. However, there has been no attempt to explain
the mechanism(s) involved in maintaining populations in spite of such
low intensive and extensive frequencies. The opportunity to study this
phenomenon was provided to me by the discovery of a reasonably large
colony of Menander felsina (Hew.) near Rio de Janeiro, Brazil. Whereas
the ecology of this butterfly resembles closely that of other forest-
dwelling but less common species, it also provides the basis for con-
clusions of a general nature applicable to other members of this group.

The purpose of this paper is to describe the adult behavior of M.
felsina in connection with feeding, male patrolling, and mating. The

38 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

results of a marking-recapture program are presented with an analysis
and discussion of the population parameters.

METHODS

The observations that form the basis of this study were gathered over
a period of more than 4 years at Pedra de Itauna, an area near the
coast west of Rio de Janeiro, Brazil. The vegetation is typical of the
restinga (coastal dune community) described in detail elsewhere (Cal-
laghan, 1977). The area for the mark-recapture study was a trail some
125 m in length just inside the south (seaward) side of the low woods
surrounding the Itauna Rock. Because of the density of the brush,
little collecting was possible elsewhere in the forest. The mark-recap-
ture project took place over a 14-week period, from 18 July-15 November
1973. Marking was done every weekend except for five near the end,
which were lost as a result of inclement weather. An average of ca. 5 h
collecting (900-1400 hrs) per day was devoted to the experiment on
study days. Individuals, once captured, received a predetermined mark
that indicated the day and were immediately released at the same spot.
Occasionally, a butterfly would show signs of shock after marking by
fluttering to the ground, in which case it was killed and eliminated
from the experiment. For analyzing the data, the Fisher and Ford
method (Fisher & Ford, 1947) was used because of the relatively small
sample sizes.

RESULTS

General Behavior: Feeding, Mating, and Patrolling

Menander felsina inhabit exclusively the low woods and nearby flats
on the seaward side of the forested areas. They prefer open habitats
such as trails and small clearings in the woods, shunning completely
the deep shade found in the higher forests. Yet, they are one of the
few butterflies that fly on cloudy days, albeit in smaller numbers than
on sunny ones. Both sexes can be found sunning in the early morning
hours with wings outspread on the upper surfaces of leaves 2-3 m
above the ground. When disturbed, they fly off with a rapid, jerky
flight, circling a few times before settling, wings outspread on the
undersides of leaves not far from where they departed. After a few
minutes they sometimes return to the same spot. When resting, they
appear to thermoregulate, raising the wings to sharper angles depending
on the intensity of the sun’s rays.

Group feeding takes place in the morning hours from 800-1100, and
to a lesser degree in the late afternoon. The M. felsina will visit prac-

VOLUME 32, NUMBER 1 39

tically any plant that happens to be in bloom in the habitat, often
becoming so “engrossed” in feeding that they can be removed with
forceps. Males and females feed together, the latter being found com-
monly only at this time.

Starting ca. 1230 hours, males begin taking up positions on the edges
of trails and clearings to await females. They sit motionless for long
periods on the upper or undersurfaces of leaves from 0.2—2 m above the
ground with their abdomens slightly raised. Males seldom leave their
perches to investigate other species of butterflies. The only ones in-
vestigated were small white pierids (Eurema sp.), which to some extent
resemble M. felsina in flight. Larger butterflies and skippers are com-
pletely ignored, which indicates that sight plays a role in the recogni-
tion of rivals or mates. This activity continues until ca. 1500. This
behavior is in contrast to the habits of lycaenids (Powell, 1968; Scott,
1974) and skippers (MacNeill, 1964). In these cases the males would
investigate any object flying past them, including small rocks (MacNeill,
1964).

To determine the extent to which spacing occurs among males, a
section of trail some 20 m long was observed on three occasions. In
the middle of this area is an opening in the woods that faces outward
toward the knee-high vegetation of the flats. Here, an older, slightly
worn male took up a station at 1255. From time to time he would
take off, flying around in an area some 5 X 3 m and perching for a few
minutes on various plants within this area. At 1326, two fresh males
moved into the study area, taking up positions to the right and left of
the older male. When one of these flew near the older male, the latter
rose up, flew around in circles with the other for a few seconds, then
returned to his original spot, the fresh male alighting a few yards out-
side the area of the older male. At 1405, a female alighted on the
outward side of the older male’s area. He immediately flew over to
the female, who took off, and followed her out onto the flats. A half
hour later, he had not returned, nor had other females appeared in the
areas of the fresh males. The next day at the same time, the same
older male was again observed in the same area.

On another occasion only fresh males were observed. They showed
less exclusiveness with regard to patrolling areas than the older males.
Another male passing nearby would sometimes be engaged in a circular
mutual chase, but both individuals would then settle down on leaves
sometimes only a meter apart. Once a male left an area, another would
often move in. During the observation periods, no females were seen
entering the fresh males’ areas.

40 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TasLe 1. Basic capture-recapture data on a population of Menander felsina.
The capture or recapture of each individual butterfly is denoted by 1.

Aug. Sept. Oct. Nov.
18 1 9 17 24 a 14 20 15
1 iL
it il
1 1
1 1
if il
if 1
il 1
Il Il 1
il 1 1
1 i
1 Il
il i
1 I
1 Ht
1 il
iL 1
Jl 1
1 iH
1 1
it it
1 1
Il il il
Hi it
il 1
1 1
a I
i il
i 1
1 1
1 Ju
1 1
1 1
1 1
1 IL
1 1
1 1
iL 1
1 1

Tatal Malaise, 122 kL none 17 ile 19 7

bo
i
bo
iN)

I conclude that M. felsina males do engage in spacing, the older
individuals defending their chosen areas more vigorously than the
younger, more inexperienced males. That the older male was successful
in meeting a female suggests that they may choose and defend the
preferred “rendezvous” areas.

VoLUME 32, NUMBER 1 Al

TABLE 2. Type A data trellis derived from data in Table 1. Units under “Date
of Marking” refer to marks and not animals.

Date of Marking

Date Captured Released 18 1 9 7) 24 ve 14 200 t5
18 Aug. 22, 22,

1 Sept. 14 14 2

9 Sept. 15 15 2 4
17 Sept. i ley i! 4 8
24 Sept. 15 15 ip I.

7 Oct. 19 19 1 2;
14 Oct. ba ha 1
20 Oct. 24 24 5 3
15 Nov. 22, 22, 2 5

1 Low captures due to inclement weather.

On one occasion, a complete courting sequence was observed. About
1416, a female alighted on a leaf, wings outspread ca. 1 m from a
perched male, who immediately flew around her several times, then
alighted and, with wings moving slowly up and down, walked to a
face to face position. There they remained for ca. 30 sec. Then, the
female walked around to the underside of the leaf, the male followed,
and copulation was initiated. This lasted for ca. 22 min. On another
occasion, a pair discovered in copula under a leaf remained so for 8
min before breaking off. Sexual activity continues until late in life.
On 15 November 1973, a male that had been marked three weeks
previously was found in copula with a freshly emerged female. Finally,
mating appears to be done by older territorial males since, of the three
cases observed, two involved males that had been previously marked.

POPULATION SIZE AND MorTALITY

Tables 1 and 2 show the basic marking and recapture data gathered
during the study, after Sheppard & Bishop (1973). It is instructive to
note that few individuals were captured more than once, which indi-
cates that collecting was not very efficient over time. Low captures on
14 October were due to rainy weather. The survival rate (Fisher &
Ford, 1947) was 0.45 per week or ca. 0.91 per day. The average life
span was 1.82 weeks by the formula E = il , which as-

1 — survival rate
sumes that all deaths occur just before sampling. Potential survival is
up to 5 weeks in the field, as confirmed by a recaptured female. Ob-
served field survival for males is up to 4 weeks. When the Lincoln
index modified by Bailey (1952) is applied to the study data, the weekly

42 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TABLE 3. Population composition over time of new and recaptured butterflies,
considering all animals captured and recaptured before and after a determined date
to constitute part of the population even though they were not captured on that date.

Date Total New % Recaptures %
18 Aug. DD, 22, 100 —- -
1 Sept. 17 WP, WL 5 29
9 Sept. 20 9 45 We 55
17 Sept. 19 4 DAML 116) 79
24 Sept. 16 13 81 3 19
7 Oct. 23 19 82 A 18
14 Oct. 11 6 54 5 46
20 Oct. DB) 16 64 9 36
15 Nov. 22, Ike 68 Tf ol
Average 19.4 10.4 = WA! =
Standard Error +22.6% +49.2% _ +42.4% ~

estimates of total population size show great variation as a result of
the large differences in recaptures from one sampling to the next.

These fluctuations were felt to be more due to inaccessibility of
individuals because of the thick brush on either side of the trail than
to changes in the population level. Therefore, to make the capture data
more realistic a third table was mounted assuming that those animals
captured in week one and recaptured in week three, for example, formed
part of the population in week two even though they were not captured
at that time. In Table 1 we see that on 18 August, 22 butterflies were
marked and released of which five were later recaptured: two on 1
September, two others on the 9th, and one on 19 September. On the
Ist, at least three butterflies were not captured.

Therefore, rather than a total of 14 for that date, we have 17 which
we know formed part of the population at that time: 12 new plus 2
recaptured plus 3 which were recaptured at later dates. The numbers
thus derived were entered in Table 3. The procedure was conducted
for each capture period during the study.

This results in considerably smoother total capture figures, which
show that if allowance is made for inefficiency in collecting, the esti-
mates of numbers of individuals in the study area was quite stable over
the 14-week period (Table 3) with a mean of 19.4 individuals and a
standard error of 4.4 or + 22.6%. More consistent results can be ob-
tained by eliminating data for 14 October. When efforts were made
to capture all the M. felsina in the study area—22 July 1973 and 6 June
1974—21 and 22 individuals were recorded, respectively, which again
suggests that the number of M. felsina in the study area remains stable
over long periods with a low number of individuals.

VoLUME 32, NUMBER 1 43

Sex ratio data gathered during the study were unsatisfactory as a
result of the similar appearance and behavior of males and females,
except during oviposition and territorial displays. Because of the delicate
nature of these butterflies, they had to be kept in the net during marking
and afterwards immediately released. This prevented detailed examina-
tion. Females, however, were a small minority of all captured. On
the two occasions referred to above, 19 males were captured each time
with 2 and 3 females, respectively. The reason is that the females
entered the study area sporadically only for mating and feeding, thus
not being as accessible as the males. The number of females caught
under these circumstances is not truly representative of the female
population; thus, for the purpose of this study, males and females were
considered together.

DIscUSSION AND CONCLUSIONS

As noted above, tropical butterfly populations, especially those of
theclids and riodinids, are characterized by low intensive and extensive
frequencies. The data from the marking-recapture program and other
observations on the M. felsina population permits a number of sugges-
tions as to why this is so.

‘Low Intensive Frequency

Low intensive frequency means that an animal is represented by a
small number of individuals in a given population. As shown by the
recapture data, the number of M. felsina frequenting the study area
remained low and fairly constant for long periods. Other students have
made similar observations on neotropical butterfly populations. Benson
& Emmel (1973) observed a roost of Marpesia berania (Hewitson) in
Costa Rica that maintained constant equilibrium of population size for
more than 3 months because of constant rates of recruitment and
mortality. Ehrlich & Gilbert (1973) recorded a similar structure for
Heliconius ethilla Godt in Trinidad over a period of 2 years, as did
Young (1972) for Siproeta epaphus (Latreille) in Costa Rica. Why do
these populations have an equilibrium level and what is the mechanism
that enables them to maintain it? Ehrlich & Gilbert (1973) explain the
constant population level in H. ethilla through the constant recruitment
of new adults over time which equals mortality. This was thought to
be due to unvarying predator pressure on the immatures and not be-
cause of a lack of foodplant, which was quite common in the habitat.
The other factor was limited adult nectar resources, which controlled
egg production, thereby regulating the production of immatures. Young

44 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

(1972) in his study of S. epaphus likewise found that egg and adult
numbers remained constant throughout the study period. As a mecha-
nism, he suggested low fecundity and low adult vagility as well as the
sheltered nature of the forest understory community. Both studies dis-
count outmigration as a factor.

My observations on M. felsina allow the following conclusions to be
made. First, neither foodplant nor nectar availability appeared to be a
limiting factor. The larval damage was always small compared with
the amount of foodplant available, showing little pressure on foodplant
resources. The adult’s preference for nearly anything that is in bloom
and its willingness to travel good distances eliminates lack of adult
nectar sources as a possibility. The factors that appear to be most
significant are low egg-laying frequency, increased longevity, and male
spacing.

Low egg-laying frequency was observed on several occasions. The
maximum number of eggs seen laid in any one afternoon by the same
female was four, these being laid singly and widely dispersed on the
foodplant (Callaghan, 1977). The results of this were shown in the
low numbers of widely separated larvae in all instars that could be
found on the same foodplant. Pupation and emergence were likewise
staggered, meaning a fairly even flow of adults over time. Since the
larvae were attended and protected by ants (Callaghan, 1977), preda-
tion was kept down to a minimum and thus did not appear to be a
significant factor as it was in the case of H. ethilla (Ehrlich & Gilbert,
1973). For M. felsina, then, the low frequency of oviposition assures
the low intensive frequency of its populations. Why egg production
should be low could not be determined during the course of the present
study.

Another factor is longevity. The 1.82 weeks observed for M. felsina
is high when compared with holarctic butterflies, but low with respect
to those tropical nymphalines that have been studied. Scott (1974)
reported an average life span of 4.2 days of Lycaena arota Bois with
a potential of 8 days. Labine (1968), Turner (1971), and Cook e¢ al.
(1971) reported the life expectancy after marking for 6 holarctic species
in the field as from 2.8-12.1 days. Powell (1968) gave a maximum of
16 days for Incisalia iroides (Boisduval). On the other hand, Benson &
Emmel (1973) demonstrated that the neotropical nymphaline Marpesia
berania has an average longevity of 43.9 days with a potential of at
least 157. Heliconids are especially long lived. Benson (1972) reported
Heliconius erato petiverana Doubleday in Costa Rica with average
observed longevities of 52 days and an individual alive 6 months after

VOLUME 32, NUMBER 1 45

marking. Turner (1971) and Ehrlich & Gilbert (1973) reported similar
results. Greater longevity ensures low intensive frequency and is of
considerable selective value in the tropics since it enables the wide-
spread dispersal of eggs in both space and time, which may diminish
parasitism and ensure a larger number of offspring reaching maturity
(Benson & Emmel, 1973). Also, having individuals in all stages of
development would permit survival of a natural disaster that might
eliminate one stage but not the others. The causes of adult mortality
of M. felsina are not precisely known, since no actual predation or other
forms of natural mortality were observed during the study. However,
potential predation exists in the form of lizards, spiders, ants, and birds.
Rapid flight and hiding beneath leaves are two methods used by adult
M. felsina and many other riodinids to avoid predation. On the other
hand, its fairly sedentary habits might mean greater predator pressure
and lower survival rate for M. felsina than for heliconines and nympha-
lines, which enjoy mimicry, distastefulness, and/or strong flight. Lower
survival would be more likely true for males because of their conspicu-
ous behavior.

Finally, male spacing seems to select for longevity, since older males
have been observed to be more aggressive and fixed to their habitual
areas and thus are easily able to drive off younger, more inexperienced
newcomers. The result is that females appearing at the rendezvous area
will be more likely to mate with strong, long-lived males. This is turn
assures longer-lived offspring, which can effectively distribute their eggs
widely in space and time. These three factors combined, then, might
provide for the perpetuation of the low intensive frequencies observed
in the M. felsina population. Although to date other forest riodinids
have not been studied in such detail, I suspect that their low intensive
frequencies may be explained on much the same basis.

Low Extensive Frequencies

Low extensive frequency means that populations are found rarely
within a given faunistic region. This is the case with M. felsina as well
as other forest riodinids. The key factor here in the case of M. felsina
is foodplant distribution. Individuals of Norantea brasiliensis Choisy
(Marcgraviaceae), the M. felsina foodplant, are widely and sparingly
distributed on the restinga, but this does not prevent their being visited
by ovipositing females.

There is a tendency on the part of the newly emerging butterflies to
establish a colony at a suitable locality near the site of their foodplant.
This was observed on several occasions, the most notable of which was

46 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

when two males were seen engaging in patrolling behavior on a small
clump of bushes near a foodplant some 500 m from the main colony.
These same two individuals were there one week later, a little bit worn
but still recognizable. Before the next visit, they had disappeared and
no others had taken their place. On the nearby foodplant, larvae were
discovered in various instars. I did not observe any other case of actual
colonization occurring on that particular clump of bushes. Undoubtedly,
this attempt failed as a result of the rather harsh, unsheltered conditions
in that particular section of the restinga. Females, then, are the col-
onizers for M. felsina, traveling considerable distances many times in
search of foodplants on which to lay their eggs.

This phenomenon has been recorded indirectly with other riodinid
butterflies. On numerous occasions a single butterfly will be encoun-
tered at a particular place and time, whereas subsequent visits under
comparable conditions will fail to turn up additional specimens. In the
same woods as the M. felsina habitat, a single male Calydna lusca
(Geyer) was captured on 18 July 1973. On 7 July 1972, a male and
female Nymphidium lisimon attenuatum Stichel were taken and a single
female Leucochimona philemon (Cramer) was taken on 7 July 1973.
Before and since, 46 collecting days over 4 years have failed to reveal
additional examples from this small wood. The best explanation is that
the females are very vagile, always on the move searching for new
foodplant localities. Similar conclusions have been reached by other
students of tropical riodinids. Ebert (1969) mentions that little species
such as riodinids “migrate continuously within a great area of favorable
biotypes. . ...”

Finally, the question arises concerning the barriers that female rio-
dinids will cross in their movement. In the case of M. felsina, open flat
areas do not appear to provide a serious obstacle, although this might
be expected because of their preference for a low forest habitat. Ob-
servations on deep forest riodinids are few but significant. On 19
January 1975, a lone female Nymphidium mantus (Cramer) was ob-
served passing through dry secondary shrub near Linhares, Espirito
Santo, Brazil, an area very different from its normal habitat in the deep
forest near the edges of swamps. Water does not appear to be a sig-
nificant barrier since many riodinid populations on either side of large
rivers such as the Amazon are virtually indistinguishable (Callaghan,
in prep. ).

SUMMARY

The results of the marking-recapture and observations of adult be-
havior allow a number of conclusions to be drawn with respect to the

VOLUME 32, NuMBER l AT

population structure of the neotropical riodinid butterfly M. felsina. It
was found that this butterfly exhibits the structure common to most
forest butterflies, that of low intensive and extensive frequencies. The
reasons for the former are low egg-laying frequency, longevity, and male
spacing. The latter was explained by a combination of high female
vagility and low intensive and extensive foodplant distribution, which
lead to the establishment of new colonies by females at widely dispersed
foodplant localities. Depending on conditions at these localities, such as
parasitism and exposure to the weather, the colony may become extinct
only later to be reestablished by another wandering female. It is sug-
gested that similar population structures for other neotropical forest
butterflies, particularly riodinids, may be explained on this same basis.

ACKNOWLEDGMENTS

I wish to thank Drs. Woodruff Benson and Keith Brown for making
many helpful comments on the manuscript and the former especially
for his helpful discussion and encouragement during the initial phases
of the field work. To Dr. Alfredo de Rego Barros my thanks for use
of the facilities of the Museu Nacional do Rio de Janeiro.

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Bates, H. W. 1864. The naturalist on the River Amazons. University of Cali-
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Benson, W. W. 1972. Natural selection for Miillerian mimicry in the Heliconius
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Benson, W. W. & T. C. Emmet. 1973. Demography of gregariously roosting
populations of the nymphaline butterfly Marpesia berania in Costa Rica.
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CaLLAGHAN, C. J. 1977. Studies on restinga butterflies. I. The life cycle and
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Crane, J. 1955. Imaginal behavior of a Trinidad butterfly Heliconius erato hydara
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EnruicH, P. R. & L. E. Gimperr. 1973. Population structure and dynamics of
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MacNeri, C. D. 1964. The skippers of the genus Hesperia in western North
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Lepidoptera by capture-recapture methods. J. Res. Lepid. 12: 135-144.
Turner, J. R. G. 1971. Experiments on the demography of tropical butterflies.
II. Longevity and home-range behaviour in Heliconius erato. Biotropica 3:

21-31.

Younc, A. M. 1972. The ecology and ethology of the tropical nymphaline but-
terfly, Victorina ephaphus. I. Life cycle and natural history. J. Lepid. Soc.
26: 155-170.

Journal of the Lepidopterists’ Society
32(1), 1978, 48

THE INTERNATIONAL CODE OF ZOOLOGICAL NOMENCLATURE

The draft third edition of the International Code of Zoological Nomenclature
is now available for comment by zoologists. Copies may be obtained (price £2.50
surface mail, £5.00 air mail) from the Publications Officer, International Trust for
Zoological Nomenclature, c/o British Museum (Natural History ), Cromwell Road,
London SW7 5BD, U.K. Comments should be sent as soon as possible, and in any
case before 30 November 1978, to the Secretary, International Commission on
Zoological Nomenclature, at the above address.

A paper explaining the major changes proposed by the Commission’s Editorial
Committee to the existing Code has been published in the Bulletin of Zoological
Nomenclature, vol. 34, part 3. Copies may be obtained (price 50p) from the same
address as copies of the draft Code.

Journal of the Lepidopterists’ Society
32(1), 1978, 49-54

AN ANALYSIS OF THE HELIOTHIDINE TYPES (NOCTUIDAE)
OF HERMAN STRECKER WITH LECTOTYPE DESIGNATIONS

D. F. HAarpwIick

Biosystematics Research Institute, Ottawa, Canada

ABSTRACT. The authenticity of the nominal type specimens of species of
Heliothidinae described by Herman Strecker is evaluated. A number of the nom-
inal type specimens are judged to be spurious. Lectotypes for the following Strecker
species are selected: Schinia approximata, Schinia dolosa, Heliothis fastidiosa,
Schinia labe, Schinia lora, and Schinia pyraloides.

In anticipation of future revisionary work on the Heliothidinae I took
the opportunity in October of 1976 to examine Herman Strecker’s type
material belonging to this subfamily at the Field Museum of Natural
History in Chicago. Strecker’s species names have always presented a
problem to Lepidopterists, firstly because his original descriptions were
often very brief, and secondly because he evidently had the habit of
augmenting or replacing his original type series. Thus, although Heliothis
regia was described from a single specimen, there are now six specimens
each labelled in his hand as “type” of regia.

In the earlier years of his career, Strecker evidently had no type con-
cept, or at least a very nebulous one. As a result, many of the specimens
on which he based his early original descriptions must have been either
destroyed or misplaced. In later years, however, with increasing aware-
ness of the value of type specimens, Strecker presumably tried to rectify
his earlier laxity by labelling specimens other than the “originals” as his
types.

If such substitutions can be demonstrated, then obviously the spurious
types have no status under the “Rules.” Nevertheless such pseudotypes
do have value in indicating Strecker’s concept of his species in, the
maturity of his later years, and should be considered in any subsequent
neotype selection procedures if these are found to be necessary. At the
present time the Strecker Collection is housed as a separate entity within
the collections of the Field Museum of Natural History, and Strecker’s
arrangement of species and his hand-printed labels have been retained.
In my discussion of type specimens which follows, the species names are
arranged alphabetically.

Rhododipsa aden Strecker

Strecker, 1898, p. 11.
The original description of aden was based on a single specimen. The male
labelled as “original type” in the Strecker collection matches the original descrip-

50 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

tion well and is evidently the one on which Strecker based his description. It is
labelled as follows: “Col.”; “384”; “S. Aden, Orig. Type’. There is a major piece
of the left hind wing broken off and the anal angle of the right front wing is
missing. A genitalic slide (no. FM Hel 1) has been prepared from the holotype.

Schinia approximata Strecker

Strecker, 1898, p. 10.

This species was described on the basis of three specimens collected by Boll
near Dallas, Texas. The three females labelled as “original types” match the orig-
inal description well and are assumed to be authentic. Because of its superior
condition the specimen numbered 76 is hereby selected as lectotype; it is labelled
as follows: “76”, “S. approximata, 374, Orig. Types”. A genitalic slide (no. FM
Hel 2) has been prepared from the lectotype.

Schinia ar Strecker

Strecker, 1898, p. 10.

The single male in the Strecker Collection labelled as “original type” matches
the original description well, and is evidently the one on which the name was
based. The specimen is labelled as follows: “371”; “S. ar, 371, Orig. Type”. A
genitalic slide (no. FM Hel 3) has been prepared from the holotype which is in
excellent condition.

Schinia dolosa Strecker

Strecker, 1898, p. 9.

The original description was based on two specimens taken near San Antonio
by Boll. The two males in the Strecker Collection labelled as “original types” are
evidently authentic. I hereby select the slightly larger specimen as lectotype; it is
labelled as follows: “tex”; “S. Dolosa, Orig. Type”. A genitalic slide (no. FM
Hel 3) has been prepared from the specimen.

Heliothis fastidiosa Strecker

Strecker, 1876, p. 121.

The original description of fastidiosa was based upon two specimens collected
by Boll in Texas. The two males in the Strecker Collection match the original
description well and undoubtedly represent the specimens on which it was based.
I hereby select the smaller specimen bearing the individual “31” label as lectotype.
The specimen is labelled as follows: “31”; “S. Fastidiosa, 31, Orig. Types”. A
genitalic slide (no. FM Hel 5) has been prepared from the lectotype.

Heliothis gloriosa Strecker
Strecker, 1877, p. 132.

A single specimen in the Strecker Collection is labelled as “original type” and
this is evidently the one on which the original description was based. The specimen,
a female, is in excellent condition except for lacking a portion of the right antenna.
It expands 1%,” and is labelled as follows: “18”; “S. gloriosa Orig. Type”. A
genitalic slide (no. FM Hel 6) has been prepared from the holotype.

Schinia hanga Strecker
Strecker, 1898, p. 9.
The species was described on the basis of one specimen collected by Boll at
Dallas, Texas. The male in the Strecker Collection labelled as “original type” is

VOLUME 32, NUMBER l 5li

evidently this specimen. It expands 144” and is labelled as follows: “70”; “393”;
“S. Hanga, Orig. Type”. A genitalic slide (no. FM Hel 7) has been prepared from
the holotype, which is in excellent condition.

Heliothis imperspicua Strecker

Strecker, 1876, p. 122.

The original description of imperspicua was based upon a single specimen, bear-
ing the number 53, which was collected in Texas by Boll. There are two specimens
in the Strecker Collection each labelled as “original type” but these are evidently
both spurious. One specimen is labelled as having been collected in Colorado;
the other is without a locality label and bears the number “49”. Neither specimen
differs from the rather generalized original description in any striking detail. The
true type of imperspicua must be presumed lost.

Heliothis inclara Strecker

Strecker, 1876, p. 122.

The original description was evidently based upon a single specimen collected
by Boll in Texas, which was numbered 46. There are now two specimens in the
Strecker Collection labelled as “original types”. One of these bears the number
78 and is considerably smaller than the specimen cited in the original description.
The other specimen is without collection number but corresponds well with the
original description and may be the true type.

Schinia labe Strecker

Strecker, 1898, p. 10.

The original description of labe was based upon two specimens collected at
Dallas, Texas by Boll. The two specimens in the Strecker Collection labelled as
“original types” are apparently these. I hereby select the smallest of the two,
which bears a separate “372” label, as lectotype. The lectotype is a male ex-
panding slightly less than 34” and is labelled as follows: “372”; “S. Labe, 372, Orig.
Types’. A genitalic slide (FM Hel 10) has been prepared from the specimen.

Heliothis lanul Strecker

Strecker, 1877, p. 132.

There is a single male in the Strecker Collection labelled as “original type” and
this is evidently the specimen on which the original description was based. There
is no locality data indicated in the original description nor on the specimen. The
holotype is labeled as follows: “85”; “S. Lanul, 85., Orig. Type”. A genitalic slide
(no. FM Hel 11) has been prepared from the type.

Schinia lora Strecker

Strecker, 1898, p. 10.

The original description of lora was based on three specimens, two from Boll
collected near Dallas, Texas, and one from Heiligbrodt at Bastrop, Texas. Only two
specimens in the Strecker Collection are labelled as “original types” and these are
apparently authentic. There is another, unlabelled specimen in the collection which
may represent the third specimen of the type series. Of the two specimens labelled
as “original types” I hereby select the specimen with the separate “73” label as
lectotype. The lectotype is a male in generally good condition which bears the
following labels “73”; “373”; “S. Lora, 373, Orig. Types”. A genitalic slide (no.
FM Hel 12) has been prepared from the lectotype.

52 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Schinia neglecta Strecker

Strecker, 1898, p. 10.

The single specimen labelled as “original type” in the Strecker Collection matches
the original description well and is evidently the one on which the name neglecta
was based. According to the original description the holotype was collected at
Loveland, Colorado. The specimen is a female, expands 1”, and bears the following
labels: “Col.”; “377”; “S. Neglecta 377., Orig. Type”. A genitalic slide (no. FM
Hel 14) has been prepared from the holotype.

Heliothis nubila Strecker

Strecker, 1876, p. 122.

Strecker’s original description of nubila was evidently based on a single specimen
taken in Texas by Boll (number 48). There are two specimens in the Strecker
Collection labelled as “original types”. Neither of these matches the original de-
scription very well, there being no red shading on the underside of the wings, and
both are numbered “72” rather than “48”. I consider these specimens to be
spurious; the three types must be presumed lost.

Schinia obscurata Strecker

Strecker, 1898, p. 10.

The single specimen labelled as original type of obscurata in the Strecker Collec-
tion is obviously the one on which the original description was based. The holotype
is in good condition except for having a notch in the left forewing. The female
specimen is labelled as follows: “St. Vincent, Pa.”; “378”; “S. Obscurata, 378, Orig.
Type’. A genitalic slide (no. FM Hel 15) has been prepared from the specimen.

Schinia pyraloides Strecker

Strecker, 1898, p. 9.

The four specimens on which the original description of pyraloides was based
are in the Strecker Collection and labelled as “original types”. The type series was
taken at Glenwood Springs, Colorado by Bruce. I hereby select the male specimen
with the individual “Col”. label as lectotype. The specimen is in generally good
condition except for lacking most of the left antenna and having a slit in the right
hind wing. The lectotype is labelled as follows: “Col.”; “S. Pyraloides, Orig. Type,
Colorado”. A genitalic slide (no. FM Hel 16) has been prepared from the specimen.

Heliothis regia Strecker

Strecker, 1876, p. 121.

So far as can be determined from the original description, the name regia was
based upon a single specimen. There are, however, six specimens in the Strecker
Collection labelled as “type”. According to the original description the holotype
was taken in Texas by Boll but none of the six nominal “types” bears a locality
label. I have compared each of the specimens with the original description and
one female matches it very well, and I construe this to be the true type. It is
labelled as follows: “S. Regia, Type”. A genitalic slide (no. FM Hel 17) has been
prepared from the holotype.

Heliothis rubiginosa Strecker
Strecker, 1876, p. 122.

Strecker’s description of rubiginosa was evidently based upon a single specimen
taken in Texas by Boll; there are now six specimens, each labelled as “Type” in the

VOLUME 32, NUMBER 1 53

Strecker Collection. None of these, however, bear the “50” label mentioned in the
original description. One of them, a male, matches the original description well and
may represent the true type but the evidence is insufficient to make a definitive
judgement.

Heliothis siren Strecker

Strecker, 1876, p. 122.

Strecker’s original description of siren was evidently based on a single specimen
collected by Boll in Texas. There are now two specimens in the Strecker Collection
labelled as “original type” but neither of these bears the “45” label mentioned in
the original description. One of the two is without number label and the other
bears an “80” label. The unnumbered specimen matches the original description
well and may represent the true type but there is no way of establishing this with
certainty.

Heliothis spectanda Strecker

Strecker, 1876, p. 122.

The original description of spectanda was based upon a single specimen taken in
Texas by Boll. When the Strecker Collection was examined, no specimen labelled
as type of spectanda was found. There was, however, a specimen in the series of
Heliothis virescens bearing the number “52” which was cited in the original de-
scription as belonging to the type. The specimen matches the original description
very well and I construe it to be the holotype. A genitalic slide (no. FM Hel 20)
has been prepared from the specimen which is a female.

Heliothis sulmala Strecker

Strecker, 1878, p. 1862.

The original description of sulmala was based upon a single male taken at
Pagosa Springs (Colorado). Strecker evidently mislaid the specimen because it
was found in 1939 in a drawer of miscellaneous moths; it lacks Strecker’s charac-
teristic type label. The specimen matches the original description very well, how-
ever, and I construe it to be the holotype. The specimen is labelled as follows:
“Heliothis Sulmala Streck., Pagosa Springs Col., (Orig. Type), McCauley, Found
(1939) in a drawer with misc. moths.” A genitalic slide (no. FM Hel 21) has been
prepared from the holotype.

Schinia tanena Strecker

Strecker, 1898, p. 10.

Strecker described tanena on the basis of a single specimen taken at Bastrop,
Texas by Heiligbrodt. There is a single male in the Strecker Collection labelled as
“type” and there is no reason to doubt its authenticity. The specimen is labelled
as follows: “tex”; “380”; “S Tanena, 380, Orig. Type”. A genitalic slide (no. FM
Hel 22) has been prepared from the holotype.

Schinia ultima Strecker

Strecker, 1876, p. 122.

The original description of ultima was evidently based upon a single specimen
taken in Texas by Boll. There are two specimens labelled as “original type” in the
Strecker Collection, but both specimens bear the number “71” rather than the “49”
indicated in the original description and both specimens differ from the original
description in several details. I do not consider either specimen to represent the
holotype, and the latter must be presumed lost.

54 JoURNAL OF THE LEPIDOPTERISTS SOCIETY

ACKNOWLEDGMENTS

I appreciate the cordial co-operation of Dr. Rupert Wenzel, Chairman
of the Department of Zoology, and Dr. Eric Smith of the Division of
Insects, during my visit to the Field Museum of Natural History.

LITERATURE CITED

SrrRECKER, H. 1876. Lepidoptera, Rhopaloceres and Heteroceres indigenous and
exotic. Number 13. Reading, Pa. P. 109-124.
1877. Id. Number 14. P. 125-134.
. 1878. Lepidoptera in Annual report of the engineers for 1878. Appendix
SS. Washington, D.C. P. 1750-1865.
. 1898. Lepidoptera, Rhopaloceres and Heteroceres, indigenous and exotic.
Supplement 1. Reading, Pa. P. 1-12, pl. 1, 2.

Journal of the Lepidopterists’ Society
32(1), 1978, 54

Letter to the editor:

A Comment on Monarchs and a “Tragedy of the Commons” in Science

When the paper by Urquhart & Urquhart appeared in this journal (1976, Vol. 30:
153-158), I sat down and wrote a letter criticizing the editorial policy of allowing
an observation to be published without providing sufficient information to allow
verification by other biologists working with Lepidoptera.

While I shared the fear that publication of the exact locale of the Mexican roost
would possibly endanger it, I felt that the authors should have at the very least
volunteered to disclose the site to responsible qualified scientists researching monarch
biology.

Subsequent events have made me regret not sending in my original comment.
Incredibly, a scientist of international reputation, Lincoln Brower, was denied the
locality information by Professor Urquhart. I do not consider such secrecy to be in
the spirit of modern science, nor necessary in this particular instance.

Anyone familiar with Brower’s body of work on the monarch would not question
his scientific stature. Anyone who has seen his environmental film on the Connecti-
cut River System cannot doubt his sensitivity to ecological problems.

We all respect the effort that Professor Urquhart has put into studying monarch
migration. That does not, however, give him territorial rights over monarch roosting
areas or free him from the scientific pesponsthality of allowing other scientists to
verify his results.

Much of the controversy and ill will which apparently has followed L. Brower’s
independent visit to the Mexican monarch roosting area might have been avoided
had the study of the monarch proceeded as unselfish science rather than a race for
glory in glossy magazines.

In the future I would hope that this journal will insist that authors be willing to
disclose their study sites to responsible colleagues.

LAWRENCE E. GILBERT
Department of Zoology
University of Texas
Austin, Texas 78712

Journal of the Lepidopterists’ Society
32(1), 1978, 55-56

ATOPOTHOURES A. BLANCHARD: A SYNONYM OF
GOYA RAGONOT (PYRALIDAE)

A. BLANCHARD
P.O. Box 20304, Houston, Texas 77025

ABSTRACT. Atopothoures ovaliger A. Blanchard becomes Gaya ovaliger (A.
Blanchard ), close to, but different from Goya stictella Hampson.

Karan and Jay Shaffer, my wife and I went collecting, 17-24 May 1977,
at the Welder Wildlife Foundation Refuge, near Sinton, Texas. Dr.
Shaffer made a special effort to collect Peoriines and was well satisfied
with the results of this trip. On their way back home the Shaffers
spent an afternoon with us at Houston, so that he could examine my
collection of Peoriines. This is when he discovered that what I had un-
fortunately described as Atopothoures ovaliger (Blanchard, 1975) should
have gone under the genus Goya Ragonot.

The male genitalia of G. ovaliger are extremely close to those of G.
stictella Hampson which is not too uncommon in Texas, but the two

Figs. 1-5. Goya: 1-4, stictella: 1, male, Welder Wildlife Refuge, Sinton, San
Patricio Co., Texas, 30 June 1975 (U.S.N.M.); 2, male genitalia of same (slide A.B.
3828); 3, male genitalia of another male, same location, same date, (slide A.B.
3827; 4, same enlarged to show gnathos. 5, ovaliger, El] Rancho Cima, Hays & Comal
cos., Texas, 29 August 1975, slide A.B. 3826 enlarged to show gnathos.

56 JoURNAL OF THE LEPIDOPTERISTS SOCIETY

species are definitely distinct. The habitus of G. ovaliger (Blanchard,
1975, Figs. 1-4) is quite different from that of G. stictella (this paper,
Fig. 1 and Shaffer, 1968, Fig. 23). The differences between their male
genitalia are not so obvious. Fig. 2 shows the genitalia of G. stictella
prepared in the conventional manner. In Fig. 3 they are prepared in
the manner favored by Shaffer (1968, page 3). Figs. 4 (stictella) and
5 (ovaliger) show the enlarged gnathos and the webs or ribs which sup-
port its apical process from beneath; this is where the most obvious
difference between the two species is to be found. Dr. Shaffer, who had
the opportunity to look at many more specimens than I had, gave me
the following information: “These ribs are provided in ovaliger with a
double row of teeth (two or three to six in each row). In stictella the
number of teeth per row varies from zero to two. Counting is com-
plicated by the fact that in both species the size of the teeth varies from
large and well developed ones to tiny, barely discernible nubbins.”

LITERATURE CITED

Biancuarp, A. 1975. A new phycitine genus and species (Pyraloidea). J. Lepid.
Soc. 29: 95-97.

SHAFFER, J. C. 1968. A revision of the Peoriinae and Anerastiinae (Auctorum) of
America north of Mexico. Bull. U.S. Nat. Mus. 280, 124 p.

Journal of the Lepidopterists’ Society
32(1), 1978, 56

PROTECTIVE BEHAVIOR IN AMPLYPTERUS GANNASCUS (SPHINGIDAE)

During August 1974, I spent about two weeks collecting Lepidoptera on the
grounds of the Inter-American Institute of Agricultural Sciences, approximately
45 km SE of San Jose, Costa Rica (near the town of Turrialba). On two separate
occasions I witnessed an interesting behavioral response in the sphingid Amplypterus
gannascus (Stoll), which was common in the area. A few gannascus would some-
times remain resting high up on the whitewashed walls of the Institute buildings
until about 1000, having been attracted to these sites by the lights on the buildings
the night before. In two cases it was possible to touch individuals by means of
tossing a multi-segmented net about twelve feet long at them. The individuals
responded to being touched by releasing their grip and sailing slowly to the
ground in a slow spiralling descent, with their wings held rigidly in a swept-back
V position. Once on the ground the moths remained passive in spite of being
nudged, and only attempted to escape after being seized by hand. The appearance
of this behavior was strikingly similar to the appearance of a dead leaf wafting
to the ground from a tree, and would seem to be a behavioral adaptation to escape
predators by imitating an unappetizing plant fragment.

Jerr Ross, Department of Museum Science, Texas Tech University, Lubbock,
Texas 79403.

Journal of the Lepidopterists’ Society
32(1), 1978, 57-58

OENEIS ALBERTA (SATYRIDAE) IN MONTANA

Oeneis alberta Elwes has been taken to date in widely scattered colonies from
Alberta, Manitoba and Saskatchewan (alberta), Colorado (oslari Skinner), Arizona
(daura (Strecker) and New Mexico (capulinensis Brown). On 19 and 20 May
1976, a series of 21 males and 6 females of alberta was taken by the author from
the high grasslands in the Little Snowy Mountains of central Montana, Fergus and
Golden Valley counties. This is a new state record for the species in Montana.
Habitat of the Golden Valley County colony is pictured in Fig. 1. Elevation is
approximately 6500 feet. Specimens from the colony are shown in Fig. 2. The
Montana colonies represent the nominate subspecies.

Additional colonies of alberta will probably be discovered in Montana as suitable
habitat in areas east of the Continental Divide is explored at the proper time of year.
Colonies should also be expected to occur in Wyoming and Utah, though none have
been found thus far.

STEVE KoHLER, Montana Department of Natural Resources and Conservation,
Division of Forestry, 2705 Spurgin Road, Missoula, Montana 59801.

Fig. 1. Habitat of Oeneis alberta in the Little Snowy Mts., Golden Valley Co.,
Mont.

58 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Mig. 2. Oeneis alberta from the Little Snowy Mts., Golden Valley Co., Mont.
(a & c) males, dorsal; (b & d) same, ventral; (e & g) females, dorsal; (f & h) same,
ventral. All photos natural size.

Journal of the Lepidopterists’ Society
32(1), 1978, 59-60

NEW OR INTERESTING LEPIDOPTERA RECORDS FROM
WESTERN TEXAS

The Panhandle and South Plains areas of Texas have probably received less close
attention from lepidopterists than other areas of the state because of the dearth of
resident collectors and the greater number of interesting species in other parts of
Texas. In view of the relative lack of information on Panhandle-Plains species, it
seems worthwhile to publish certain significant records from my collection and the
Texas Tech University collection at this time. I report herein one new Texas record
(Pieridae) and additional records of species not usually associated with these areas
of Texas.

PIERIDAE

Kricogonia lyside (Godart) has previously been reported from the Panhandle-
Plains area only in October (Kendall and Freeman 1963, The Butterflies and Skippers
of Texas: A Tentative List, Sinton, Texas, 6 p.). The Texas Tech University col-
lection contains two males of this species from Lubbock, Texas (Lubbock Co.),
both of which are in very good condition. One was collected on 12 July 1970 by
D. W. Kiser, and the other on 24 September 1967 by U. Barber. In addition to
these specimens, I collected one male and two females in fair condition at Palo
Duro Canyon State Park, Randall Co., Texas, on 6 May 1977.

Phoebis agarithe maxima (Neumoegen). Although this species was not cited in
Kendall and Freeman’s checklist as having been recorded from the Panhandle-Plains
region of Texas, the Texas Tech collection contains two males, in fair condition,
from Lubbock, Texas (Lubbock Co.). One was collected on 7 July 1967 by
“E J W,” and the other on 16 September 1970 by P. M. Allen. I observed numer-
ous males and females of agarithe maxima in Lubbock, Texas throughout September
of 1976, and captured a single worn male on 19 September 1976.

Pieris napi (Linnaeus). The Texas Tech collection contains a single perfect male
specimen of an undetermined subspecies collected on 17 August 1970, at Canyon,
Texas (Randall Co.) by Walt Fournier, a former Tech graduate student. As far
as can be determined, this record is a new one for the state of Texas.

LYCAENIDAE

Lycaeides melissa melissa (Edwards). I collected a single perfect male of this
species on 31 August 1975 at the Buffalo Springs Lake Recreation Area (4 mi. E
Lubbock, Lubbock Co., Texas). This record tends to support the contention by
Rickard and Vernon (1975, J. Lepid. Soc.: 150) ihat this heretofore rarely reported
species has probably just been overlooked in the past.

NYMPHALIDAE

Chlosyne janais (Drury). A single female of this common neotropical species
was collected by me at my residence in Lubbock, Texas, on 11 June 1977. The
specimen has badly torn hindwings but is in fair condition otherwise. This species
has not previously been reported from the Panhandle-Plains region of Texas.

SATURNIIDAE

Hemileuca hera hera (Warris). Although Douglas C. Ferguson states that hera
is “widespread in the West but not known from Texas” (1972, Bombycoidea-
Saturniidae in part, p. 106. in R. B. Dominick, et al., The Moths of America
north of Mexico, Fascicle 20.2B), the Texas Tech collection contains a single male
in very good condition, collected on 16 September 1969 in Dickens, Texas (Dickens
Co.) by M. Hughes.

Callosamia promethea (Drury). The Texas Tech collection contains one female,

60 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

in fair condition, collected on 2 September 1973 at Junction, Texas (Kimble Co.)
by Tech graduate student Sandy M. Benbow. According to Ferguson (p. 235),
promethea has not been cited previously as occurring west of Tyler, San Jacinto,
and Montgomery counties in eastern Texas. Thus this record suggests a possible
range extension of several hundred miles into the Edwards Plateau area. Texas
Tech University students and faculty collect annually at a field campus in Junction,
so it should be possible to determine if promethea is more than just a stray in the
area.

ACKNOWLEDGMENTS

I wish to thank Roy O. Kendall of San Antonio, Texas, for reviewing this paper
and confirming the identification of these specimens, and Dr. David E. Foster of
the Texas Tech entomology faculty for allowing me to examine material in the Tech
entomology collection.

Jerr Ross, Department of Entomology, Texas Tech Uniwersity, Lubbock, Texas
79403.

Journal of the Lepidopterists’ Society
32(1), 1978, 61-62

ERYNNIS BRIZO LACUSTRA AND HESPERIA COLUMBIA IN THE
SIERRA NEVADA

Burns (1964, U. C. Publ. Entomol. 37: 1-214) reports no records for Erynnis
brizo lacustra Wright for the Sierra Nevada, and MacNeill (1964, U. C. Publ.
Entomol. 35: 1-221) lists no records for Hesperia columbia Scudder from there
except one female in the AMNH from “Sier. Nev.” Both are indicator species of
the coast range serpentine belts north of San Francisco. Until recently, serpentine
outcrops have been little collected in the western foothills of the Sierra Nevada of
east-central California. Table 1 (next page) lists the new distribution records there.

Sometimes the adults may fly a few miles from their serpentine areas to hilltop:
e.g., both hilltop on Rocky Ridge, 1700-1900’, N. of Monticello Dam, Yolo Co., a
non-serpentine area composed of Upper Cretaceous marine rocks of the Venado
Formation. The nearest serpentine occurs in the extensive Mesozoic ultrabasic in-
trusive rocks and the Franciscan Formation some 6 miles to the west. Similarly,
Footman Ridge, Mariposa Co., is Paleozoic marine (also the area to the N & E),
and to the south is Mesozoic granitic rocks, with no serpentine nearby. The nearest
serpentine is found 5 mi. W. as Jurassic-Triassic metavolcanic rocks and 8 mi. SW
near Mariposa as Mesozoic ultrabasic intrusive rocks. In the meadows, forests, and
canyon immediately adjacent to Footman Ridge on the W & N, neither species has
ever been collected.

On 15 May 1970, E. slope Walker Ridge along Brim Grade, c. 1800’, SW of
Leesville, Colusa-Lake Co. line, I noticed a female lacustra ovipositing on the
terminal growth of a Quercus durata Jepson bush growing on serpentine soil along
a roadbank, at 1125. Bums (1964) says “the skipper invariably occurs in direct
association with QO. durata, a serpentine obligate” (see Whittaker et al., 1954,
Ecology 35: 258-288). However, in some areas, it may also use Quercus dumosa
Nutt. which hybridizes with Q. durata and grows in strictly non-serpentine soils
(see Forde & Faris, 1962, Evolution 16: 338-347 ).

Heretofore, these skippers were considered more coastal in their California
distribution.

OaxkLEy SHIELDS, Department of Entomology, University of California, Davis,
California 95616.

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JoURNAL OF THE LEPIDOPTERISTS SOCIETY

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Journal of the Lepidopterists’ Society
32(1), 1978, 63-64

BOOK REVIEWS

THE BUTTERFLIES AND Morus oF HAMPSHIRE AND THE ISLE OF WicHT (being an
account of the whole of the Lepidoptera) by B. Goater. 1974. E. W. Classey,
Ltd., Faringdon, Oxon, England. xiv + 439 pp. Price: £6.50, postpaid.

Perhaps nothing so authoritative as this book has been written on so small an
areas fauna. The book is annotated with published records, manuscript notes and
personal observations on all of the butterflies and moths of Hampshire and the Isle
of Wight since the beginning of collecting there. Make no mistake, this book is an
historical document, and as a record of what is (and was) in the County, and on
the island, it is invaluable.

Goater has drawn records from many contemporary sources, and there are some
prominent English entomological contributors to the list: such as D. W. Ffennell,
John Heath, E. C. Pelham-Clinton, and the Baron C. G. M. de Worms, names
well-known in English lepidopterology. This information provides a regional list
unlike any we have seen and the coverage is complete through 1972.

Despite the lovely picture of Argynnis paphia on the cover, this is no “coffee
table” book. It sticks strictly to business, and those looking for pretty pictures by
which to identify British Lepidoptera should be forewarned to stay away from it.
As a book of information (isn’t that what we really need? ), it is superb, and from it
one can discover when, where and at what time any species of butterfly or moth has
been captured in that area, and, if it has been reared, on what foodplant. I suspect
that in this alone the book has fulfilled its purpose, and, additionally, it should
stimulate the collector in the area to “fill in the blanks”.

The nomenclature used is standard perhaps only to the British, since it is derived
from the Kloet and Hincks Checklist of British Insects, part 2, Lepidoptera, 1972.
In this treatment, the Hesperioidea and Papilionoidea directly follow the Ptero-
phoroidea and precede the Bombycoidea. To a North American rhopalocerist this
arrangement will seem strange, even incomprehensible, since there are few other
classifications that follow this one. Most schemes place the butterflies and skippers
above the Noctuidae, the “top” family in Goater’s system. If you are interested in
the butterflies, by the way, look on pp. 214-245. The sphingids may be found on
pp. 307-312, and the saturiid (there is only one) on p. 248, while Catocala are
on pp. 404-406. This gives a bit of a “road map” to the reader just trying the book
for the first time (I confess to a great deal of initial confusion ).

As stated before, don’t buy this book on the basis of the pretty picture on its
cover. Neither is this an identification manual. But if you are interested in a superb
compendium of what is known about a limited fauna, by all means get the volume.
It sets a fine standard, despite a few typographical errors not alluded to here (they
happen to everyone! ), for future lists on small faunas.

Lee D. Miter, Allyn Museum of Entomology, 3701 Bay Shore Road, Sarasota,
Florida 33580.

BUTTERFLIES OF West MALAYSIA AND SINGAPORE by W. A. Fleming. 1975. E. W.
Classey Ltd., Farington, Oxon, England. Vol. 1: vii-x+64 pp., pls. 1-54; vol. 2:
vii-x + 92 pp., pls. 55-90. Price: £19.50, postpaid.

This book, effective as it is, is something of an enigma. I find it impossible to
rationalize making it in two volumes if the series is only to be sold as a whole, and
not broken into separate volumes, if the buyer so desires. The text is identical in
both volumes to page 15, so it is only in the plates and the parts following them
that the two volumes differ. A little elementary arithmetic shows that if the volumes

64 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

were combined into a single one, there would be a very manageable and useful
single book of four prefatory pages, 144 text pages and 90 plates. Surely this would
have been a better plan. Maybe others will have the problem I did—I would refer
to “the” book to identify a Malaysian butterfly, and I almost inevitably selected the
wrong volume. The only excuse I can think of for publishing this work in two
volumes is economic, and perhaps £19.50 for a single book would put off some
buyers, but with the price of books what it is today, I doubt it.

But enough of the complaining. There is a multitude of information in this book,
even though the style is such that it takes some acclimation. The nomenclature is
up-to-date and applied to the right insects. Species in which the illustrations are
not enough for identification are characterized in the text, and in those instances
where genitalic dissection is necessary for final determination, the fact is noted,
even though the genitalia are not figured. If Fleming had included some biblio-
graphic citations to these problem areas, and to the many included foodplants
records, the book would have been more authoritative, and the space these refer-
ences would have added could not have been that much.

The illustrations, however, are where the books truly excel. All of the photographs
of specimens illustrate the salient points well and facilitate the identification of the
insects in question. Al] of the specimens used are not perfect; some are downright
tatty, such as the illustrated female of S12, Lethe europa malaya Corbet on Plate
24, but these were the best specimens available in collecions, and the photographs
mercifully have not been “prettied up”. The color fidelity is very high, and at
least most of the specimens are fresh, rather than century-old museum relics. Iden-
tification of even the difficult Malaysian lycaenids is facilitated by them, though of
course it is not made simple—no book could achieve that!

I particularly appreciated the accurate citation of the authors of various taxa, even
though these names were not bracketed where appropriate. At long last, both of
the Felders are cited as the authors of names proposed in the “Reise Novara’, not
just a blanket “Felder”. This latter practice seems to have dated from “Seitz” where
only Cajetan Felder was given credit for the descriptions in the work, even though the
authors themselves cited “nobis” on every new name, rather than the singular “mihi”.

On balance, this is an exceilent book, the foregoing criticisms notwithstanding,
and one that is remarkably free of typographical errors. The text portions are per-
haps a bit too abbreviated, and authority is not given for many statements. I
personally would have preferred a single volume about the size of Corbett and
Pendlebury’s The Butterflies of the Malay Peninsula, but the present book accom-
plishes some things that the earlier authors could not: Fleming has made the
identification of Malaysian butterflies considerably easier than before. No more can
be asked of any author! If your interests lie in the butterflies of southeastern Asia,
by all means buy this book.

Lee D. Mitier, Allyn Museum of Entomology, 3701 Bay Shore Road, Sarasota,
Florida 33580.

EDITORIAL STAFF OF THE JOURNAL
Austin P. Puatr, Editor

Department of Biological Sciences
University of Maryland Baltimore County, 5401 Wilkens Avenue
Catonsville, Maryland 21228 U.S.A.
Dovuctas C. Frercuson, Associate Editor THEODORE D. SARGENT, Associate Editor

NOTICE TO CONTRIBUTORS

Contributions to the Journal may deal with any aspect of the collection and study
of Lepidoptera. Contributors should prepare manuscripts according to the following
instructions.

Abstract: A brief abstract should precede the text of all articles.

Text: Manuscripts should be submitted in duplicate, and must be typewritten,
entirely double-spaced, employing wide margins, on one side only of white, 8% x 11
inch paper. Titles should be explicit and descriptive of the article’s content, including
the family name of the subject, but must be kept as short as possible. The first men-
tion of a plant or animal in the text should include the full scientific name, with
authors of zoological names. Insect measurements should be given in metric units;
times should be given in terms of the 24-hour clock (e.g. 0930, not 9:30 AM).
Underline only where italics are intended. References to footnotes should be num-
bered consecutively, and the footnotes typed on a separate sheet.

Literature Cited: References in the text of articles should be given as, Sheppard
(1959) or (Sheppard, 1959, 196la, 1961b) and all must be listed alphabetically
under the heading LrreRATuRE Crrep, in the following format:

SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd. ed. Hutchinson,
London. 209 p.

196la. Some contributions to population genetics resulting from the
study of the Lepidoptera. Adv. Genet. 10: 165-216.

In the case of general notes, references should be given in the text as, Sheppard
(1961, Adv. Genet. 10: 165-216) or (Sheppard 1961, Sym. Roy. Entomol. Soc.
London 1: 23-30).

Illustrations: All photographs and drawings should be mounted on stiff, white
backing, arranged in the desired format, allowing (with particular regard to lettering )
for reduction to their final width (usually 4% inches). Illustrations larger than 8%
x 11 inches are not acceptable and should be reduced photographically to that size
or smaller. The author’s name, figure numbers as cited in the text, and an indication
of the article’s title should be printed on the back of each mounted plate, Figures,
both line drawings and halftones (photographs), should be numbered consecutively
in Arabic numerals. The term “plate” should not be employed. Figure legends must
be typewritten, double-spaced, on a separate sheet (not attached to the illustrations),
headed ExpLANATION OF FicuRES, with a separate paragraph devoted to each page of
illustrations.

Tables: Tables should be numbered consecutively in Arabic numerals. Headings
for tables should not be capitalized. Tabular material should be kept to a minimum
and must be typed on separate sheets, and placed following the main text, with the
approximate desired position indicated in the text. Vertical rules should be avoided.

Proofs: The edited manuscript and galley proofs will be mailed to the author for
correction of printer’s errors. Excessive author’s changes at this time will be charged
to authors at the rate of 75¢ per line. A purchase order for reprints will accompany
the proofs.

Correspondence: Address all matters relating to the Journal to the editor. Short
manuscripts such as new state records, current events, and notices should be sent to
the editor of the News: Jo Brewer, 257 Common Street, Dedham, Massachusetts
02026 U.S.A.

ALLEN PRESS, INC. REUSED LAWRENCE, KANSAS
Us he

CONTENTS

Types OF PARNASSIUS CLODIUS GALLATINUS (PAPILIONIDAE). Steve

Bove oO Ses PS CON La aaa i 1
SPECIFICITY, GEOGRAPHIC DISTRIBUTIONS, AND FOODPLANT DIVERSITY
IN Four CALLOPHRYS (MITOURA) (LYCAENIDAE). Kurt Johnson 3
FooppLANT, Hapirat, AND RANGE OF CELASTRINA EBENINA (LyY-
CAENIDAE). Warren H. Wagner, Jr. and T. Lawrence Mellichamp 20
Strupies ON Restinca Butrerruises. IJ. NoTes ON THE POPULATION
STRUCTURE OF MENANDER FELSINA (RiopINDAE). Curtis J.
Callaghan, 2 37
AN ANALYSIS OF THE HELIOTHIDINE TyPEs (NocrumAE) OF HERMAN
STRECKER WITH LEcToTyPE DesicNaATIoNs. D.F. Hardwick... 49
ATOPOTHOURES A. BLANCHARD: A SYNONYM OF GOYA RAGONOT
(Pyratmar). A. Blanchard 0. ee 50
GENERAL NOTES
Protective behavior in Amplypterus gannascus (Sphingidae). Jeff Robb 56
Oeneis alberta (Satyridae) in Montana. Steve Kohler __.. 57
New or interesting Lepidoptera records from Western Texas. Jeff Robb 59
Erynnis brizo lacustra and Hesperia columbia in the Sierra Nevada.
Oakley Shields i000 ee 61
Noes) AND News 2220230 ies a 19, 48
Book (REVIEWS. i eT aa a 63

Volume 32 1978 Number 2

JOURNAL

of the

LEPIDOPTERISTS’ SOCIETY

Published quarterly by THE LEPIDOPTERISTS’ SOCIETY

Publié par LA SOCIETE DES LEPIDOPTERISTES
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN
Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS

19 July 1978

THE LEPIDOPTERISTS’ SOCIETY

EXECUTIVE COUNCIL

J. W. TuwpeNn, President KENELM W. Puut.ip, Vice President
I. F. B. Common, Ist Vice President JuLtian P. DoNnAHUE, Secretary
Lionet Hiccrs, Vice President RONALD LEUSCHNER, Treasurer

Members at large:

F. S. CHEw R. A. ARNOLD J. F. EMMEL
D. F. Harpwick E. D. CasHAaTT R. R. GATRELLE
J. B. ZrEcLER R. E. STANFORD A, Po Pian

The object of the Lepidopterists’ Society, which was formed in May, 1947 and
formally constituted in December, 1950, is “to promote the science of lepidopterology
in all its branches, . .. . to issue a periodical and other publications on Lepidoptera,
to facilitate the exchange of specimens and ideas by both the professional worker and
the amateur in the field; to secure cooperation in all measures” directed towards —
these aims.

Membership in the Society is open to all persons interested in the study of
Lepidoptera. All members receive the Journal and the News of the Lepidopterists’ —
Society. Institutions may subscribe to the Journal but may not become members.
Prospective members should send to the Treasurer full dues for the current year,
together with their full name, address, and special lepidopterological interests. In
alternate years a list of members of the Society is issued, with addresses and special
interests. There are four numbers in each volume of the Journal, scheduled for
February, May, August and November, and six numbers of the News each year.

Active members—annual dues $13.00
Student members—annual dues $10.00
Sustaining members—annual dues $20.00
Life members—single sum $250.00
Institutional subscriptions—annual $18.00

Send remittances, payable to The Lepidopterists’ Society, and address changes to:
Ronald Leuschner, 1900 John St., Manhattan Beach, California 90266 U.S.A.

Back issues of the Journal of the Lepidopterists’ Society, the Commemorative
Volume, and recent issues of the NEWS are available from the Assistant Treasurer.

The Journal is $13 per volume, the Commemorative Volume, $6; and the NEWS,
$.25 per issue.

Order: Mail to Charles V. Covell, Jr.. Memoirs Editor, Department of Biology, Uni-
versity of Louisville, Louisville, KY 40208, U.S.A.

The Lepidopterists’ Society is a non-profit, scientific organization. The known
office of publication is 1041 New Hampshire St., Lawrence, Kansas 66044. Second
class postage paid at Lawrence, Kansas, U.S.A. 66044.

Cover illustration: Dasychira dorsipennata larva, dorsal and lateral views. From
Fascicle 22.2, “Lymantriidae,” by Douglas C. Ferguson, in Moths of America North
of Mexico. The drawing was done by E. R. Hodges, Scientific Illustrator, Department
of Entomology, Smithsonian Institution. (Reproduced by permission of the author. )

J OURNAL OF

Tue LeprpopreRists’ SOCIETY

Volume 32 1978 Number 2

Journal of the Lepidopterists’ Society
32(2), 1978, 65-74

STUDIES ON THE INTERACTIONS OF MORPHO PELEIDES
(MORPHIDAE) WITH LEGUMINOSAE

ALLEN M. YouNG

Invertebrate Division, Milwaukee Public Museum
Milwaukee, Wisconsin 53233

ABSTRACT. The butterfly Morpho peleides Kollar is a widespread species
throughout tropical America, exploiting several wild genera and species of Leguminosae
as larval foodplants. Field studies show that this species feeds on a broad spectrum of
wild legumes on a regional basis. This interaction was explored in the laboratory by
rearing caterpillars on peanut plants and alfalfa, cultivated legumes. The life cycle is
completed successfully on these artificial foodplants, but feeding on alfalfa taken
from an expressway led to mass mortality of caterpillars. Apparently the alfalfa was
contaminated from some environmental source. In the native habitats of this butterfly,
the Leguminosae are both diverse and numerous locally. This suggests that the mo-
nophagous feeding habit provides sufficient ecological flexibility for exploiting different
genera and species of the family. This is sufficient to maintain breeding populations of
M. peleides in secondary habitats. Forest-dwelling species of Morpho are predicted
to be experiencing different types of selection pressures favoring polyphagous feeding.

In the premontane tropical wet forest life zone (Tosi 1969) of north-
eastern Costa Rica, a larval foodplant of the butterfly Morpho peleides
Kollar (Lepidoptera: Morphidae) is the vine Machaerium aff. flori-
bundum Benth. (Leguminosae). The vine and butterfly occur in stands
of mixed primary and secondary tropical wet forest (Fig. 1). It is known
that M. peleides utilizes several leguminous woody vines and trees as
larval foodplants in Costa Rica (Young and Muyshondt 1973) and the
species can be reared on commercially available peanut plants both in
Costa Rica and Wisconsin (Young 1974). This present paper examines
further the feeding habits of M. peleides larvae, using eggs obtained
from a population in premontane tropical wet forest (rather than from
montane forest, as in a previous study), and involves plants not studied
previously (Young 1974). The results further support the assumption

66 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Mig. 1. Above: mixed primary-secondary tropical wet forest habitat of the butterfly
Morpho peleides at Finca El Tigre, near La Virgen, Heredia Province (Sarapiqui
region), Costa Rica. At the spot shown here, M. peleides and M. granadensis are
microsympatric. Below: Machaerium aff. floribundum, a forest leguminous vine
which is a larval food plant of M. peleides (and probably M. granadensis) in the wild.

VOLUME 32, NUMBER 2 67

that larvae of M. peleides are monophagous feeders on many temperate
and tropical genera of Leguminosae.

METHODS

A female of M. peleides was captured on bait of rotting bananas placed
near the edge of a forest habitat (Fig. 1) on February 14, 1977 at “Finca
El] Tigre,’ a farm adjacent to “Finca La Tirimbina,” a few km from La
Virgen, Heredia Province (Sarapiqui region), Costa Rica (220 m elev.).
She was placed in a clear plastic bag containing a fresh cutting of M. aff.
floribundum, and within eight days had produced a total of 40 viable
eggs. The female was then preserved, and the eggs were brought to
Milwaukee, Wisconsin for rearing. The eggs began to hatch on February
25 in Costa Rica and by the time the morphos arrived in Wisconsin, they
were all Ist instar. In Costa Rica these larvae were fed leaves of Dioclea
wilsoni (Leguminosae) but they were switched to peanuts (Arachis
hypogea L.—Leguminosae) upon arrival at the Milwaukee Public
Museum. The larvae were kept on potted peanut plants placed in a
covered glass tank in a laboratory. A growth light was kept over this
rearing chamber. The rearing program in Wisconsin extended from
March 3 through May 25, 1977 (the date of the late eclosion). Records
were kept on body lengths and head capsule widths of all caterpillars.
The sources of peanut plants used were (1) Olds Seeds from Madison,
Wisconsin and (2) Crop Science Department of North Carolina State
University (Raleigh). Near the end of the experiment (April 22), the
foodplant was switched to alfalfa (Medicago sativa L—Leguminosae);
at this time most of the larvae were in the late 4th instar. The alfalfa
plants used were obtained from a farm in Waukesha County, Wisconsin.
Later (May 2) the remaining 5th instar larvae (several had pupated)
were fed alfalfa collected from the side of an expressway in downtown
Milwaukee. Like the peanuts, the alfalfa plants were potted, but this
time soil brought in with the plants from the field was used. One 4th
instar caterpillar was offered a seedling of Erythrina crista-galli L.
(Leguminosae) from Brazil. Records were kept on larval survival
throughout the study. The adults obtained were kept for further
examination.

RESULTS

Both young and older larvae of M. peleides ted successfully on peanut
and alfalfa leaves in the laboratory, followed by normal eclosion (Fig. 2).
In addition, at least the 4th and 5th instar larvae will feed on Erythrina.
Although some caterpillars feed intermittently throughout the day, the

68 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fig. 2. Right column: second and fourth instar catepillars of M. peleides on peanut
leaves at the Milwaukee Public Museum; note eaten areas of leaf in first illustration.
Leaf column: fifth instar caterpillar on peanut plant and freshly-enclosed adult
clinging to empty pupa case (at Milwaukee Public Museum ).

VOLUME 32, NUMBER 2 69

MEAN AND RANGE OF BODY LENGTH (MM) OF CATERPILLARS

20

3 . .
16

14

tt

ae

MAR. 8 MAY 2
(DAY 1) SUCCESSIVE DAYS OF OBSERVATION (DAY 38)

Fig. 3. Growth and size (body length) patterns for Morpho caterpillars reared in
the laboratory. The vertical lines give the range in body lengths.

greatest amount of feeding occurred in the late afternoon and early
morning (e.g., 16:00-19:30 hrs/C.S.T.). Fourth and 5th instar larvae
rested on the rims of the pots containing the peanut plants, and they
would crawl up the plants to feed. Younger ones rested on leaves and
shoots.

Survival both on peanuts and on “farm alfalfa” was 100%. However,
larvae fed “expressway alfalfa” showed considerable mortality: between
May 4 and May 12, the number of healthy caterpillars dropped from 32
to 13. Very shortly after being fed the expressway alfalfa, many died.
Death was preceded by a drastic contraction of body length, and spasmic

70 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TasLe 1. Head capsule size statistics for caterpillars of the tropical butterfly
Morpho peleides.

No. Mean head capsule Range of head
Instar measured width (x = S.D.) capsule widths (mm )
1 17 1.50 + 0.08 1.4-1.5
2, 34 2.19 + 0.09 2.0—2.3
3 315} BA ae Vpllc 3.3-3.8
4 oll 4,91 +0.11 A4.7-5.1
5 Ia 6.09 + 0.45 5.2-6.7

Note—data obtained from molted head capsules at the end of each instar [distorted (crushed)
head capsules are not included in the calculations]. All head capsules were collected within 24 hours
after each molt.

waves of movements lengthwise. Afflicted caterpillars fell from the
plants and wriggled on the bottom of the cage before dying. When
attempts were made to replace them on the foodplant, they again lost
hold and fell off. Their feces had a reddish-orange component that was
quickly absorbed on paper toweling. A few died as prepupae.

Initial signs of the affliction included a larva remaining stationary on
the plant, with the anterior half of the body hanging off to one side.
Such a state lasted a few days before the larva fell off the plant and died.
As a result of this sudden and epidemic-like mortality, only those indi-
viduals that had already pupated by May 2 survived to emerge as adults.
A total of 12 adults, all males, were obtained. Thus, the mortality was
100% among the 19 individuals that were still larvae on May 4.

Based on a sample of ten randomly chosen larvae, molting is not syn-
chronous. The 5th instar is the longest, and despite changes in the —
foodplant type, development proceeds without major accelerations or
decelerations in the daily pattern of growth (Fig. 3). With the exception
of the Ist instar, there is considerable variation in the body length and
the magnitude of this variation is about the same for the four instars
(Table 1). Excluding a small bias introduced by unequal sample sizes,
the range of variation in head capsule width is very low for all instars
(Table 1). Body length and head capsule width are used here as estimates
of larval size. The total development time of 91 days is broken down
as follows: (1) egg = 11 days; (2) caterpillar = 65 days; (3) pupa = 15
days. The right forewing length for the adult male butterflies ranged
from 55 to 65 mm (N = 12), but the exclusion of two individuals reduced
this range to 61-65 mm.

DIscUSSION

The data are useful for discussing (1) feeding behavior of the cater-
pillars of M. peleides, and (2) apparent effects of environmental con-

VoLUME 32, NUMBER 2 fol

tamination of a foodplant of an exotic butterfly. The latter was an
unexpected outcome of the study.

Morpho accepts peanuts and other legumes, not used as foodplants in
the wild, as discussed previously (Young 1974). But to these records I
add alfalfa and Erythrina as acceptable foodplants of M. peleides cater-
pillars in the laboratory. Erythrina is native to the New World tropics
(Bailey 1969), although it is not known if it is a natural foodplant of
Morpho. Greenhouse cultures of this plant are usually infested with
herbivorous insects, suggesting few effective defenses operative against
such attacks. Alfalfa, a near relative of peanut, is commonly cultivated,
and occurs as a weed species along roadsides; it is native to the Old
World. As a weed species, alfalfa may possess few defenses against
herbivores as energy allocation is likely to be directed toward high repro-
ductive potential (Lewontin 1965). Thus, cultivars such as peanut and
alfalfa, weeds such as alfalfa, and ornamentals, such as Erythrina, are
examples of leguminous plant species with few defenses against her-
bivores, perhaps making them ideal to serve as food for Morpho larvae.
It is not known if Morpho will oviposit on these plants in the laboratory.

I observed previously that caterpillars of M. peleides are primarily
“dawn-dusk” feeders in the wild (Young 1972a), and this is also true
for laboratory cultures experiencing the Wisconsin dawn-dusk cycle
(Young 1974; pers. obs.). Apparently the larvae are programmed with
a rhythmicity for peak periods of feeding activity in both tropical and
temperate situations. In the wild, 4th and 5th instar caterpillars rest on
the trunks of the foodplant where they blend in with the background
(Young 1972a), and this behavior explains why they rested on the rims of
the pots containing the peanut plants.

Using a larger sample size, Young (1974) estimated the total develop-
mental time for M. peleides on peanuts to be about 105 days, or about
14 days longer than the estimate obtained in the present study. Body
length of 5th instars in the previous study, was about 73 mm as com-
pared to 70-71 mm in the present study. Several factors may be relevant
here: (1) the eggs in the two studies came from different regions. Thus
selection pressures could have been different in terms of effects on
development time; (2) differences in the foodplant as related to time
of the year, and other factors. The discrepancy in the development time
is in the length of the larval stage; the egg and pupal stages are the
same (Young 1974; pers. obs.). Thus, differences in the foodplant may
be involved. Switching to alfalfa (not done in the previous study ) might
have accelerated development. To test this, eggs will have to be reared
entirely on this plant in a future study. The transfer to alfalfa was done

—~l
bo

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

in the fifth instar, the time of greatest food intake. In the previous study,
older peanut plants were used, and it may well be that older plants have
resistant properties more expressed than younger ones. There is some
evidence that the defense systems of peanut plants change with age: in
the present study I observed that Morpho larvae refuse to eat the first
set of leaves of a peanut seedling, eating only shoot and leaf tissue above
this point (S. Borkin and A. Young, pers. obs.).

The observed high level of mortality among 5th instar larvae on alfalfa
plants taken from the downtown expressway may have been due to a
contamination of these particular plants by an industrial or automobile
residue. An exotic insect such as Morpho, when exposed to a con-
taminated foodplant, may be expected to encounter such mortality.
Since only male adults were reared I presume that all the females died as
Sth instar larvae, since M. peleides has a sex ratio of unity (Young 1972b,
1973; Young and Muyshondt 1973). Female larval development generally
takes longer than that of males. The observed inability of afflicted larvae
to grasp the foodplant and feed was very likely due to the contaminant
affecting the nervous system. It is not likely that a strain difference
affecting feeding ability by morphos exists between the expressway
alfalfa and farm alfalfa, since larvae did eat the former and until the time
of obvious signs of illness, their feeding behavior appeared normal.

The data indicate that feeding flexibility of M. peleides caterpillars is
considerable in the sense that it allows this monophagous tropical her-
bivore to exploit a broad range of genera and species locally. Foodplant
records for M. peleides from Costa Rica and El Salvador indicate that
many different wild Leguminosae are used, and allied South American
species exhibit similar behavioral flexibility (Otero 1971; pers. comm. ).
Secondary forest habitats in the wetter regions of Central America locally
support a wealth of leguminous vine, shrub, and tree species, many of
which are used by M. peleides. It is therefore not surprising that cater-
pillars will feed successfully on allied legumes not used as natural food-
plants, including cultivated forms such as peanuts and alfalfa. On a
per unit area basis, secondary habitats in the tropics support large
patches of Mucuna, Dioclea, Machaerium, Inga, etc. and many patches
may occur locally. Thus, in terms of larval foodplants, the environment
is very certain or predictable creating selection pressures favoring eco-
logical specialization such as monophagy. As foodplant patches increase
in number and size in an area, monophagy is considered an optimal
feeding strategy for a herbivore (Levins and MacArthur 1969).

Some species of Morpho that live in the canopy of primary tropical
forest deposit eggs on different families of trees and woody vines (Otero,

VoLUME 32, NUMBER 2 73

pers. comm.) and these species may be polyphagous. Miller (1968) lists
several Morpho foodplant families, although the degree to which each
species oviposits on more than one family has not been determined.
Polyphagy in Morpho is adaptive in habitats where individuals of each
foodplant species are greatly dispersed over large areas, making it ener-
getically difficult to exploit a single family of foodplants. In forest hab-
itats where each foodplant species is greatly dispersed over large areas, the
environment is less certain in terms of a female butterfly locating suc-
cessfully an individual of that particular plant. As more plants are added
to the local foodplant niche, the environment becomes more certain; the
incorporation of additional local foodplants implies the evolution of
polyphagy, since member genera and species of individual plant families
in tropical forests are greatly dispersed over large areas. Thus, although
M. peleides and its near allies such as M. achilles may exhibit considerable
feeding flexibility within the Leguminosae, they are monophagous species;
polyphagous species of Morpho are expected to occur in primary forests.
These include such likely candidates as M. amathonte, theseus, grana-
densis, and cypris in the Central American rain forests.

ACKNOWLEDGMENTS

This research would not have been possible without the financial
support of the Friends of the Museum (of the Milwaukee Public
Museum) and James R. Neidhoefer. Susan Borkin and Joan Jass ( Mil-
waukee Public Museum) conducted the rearing studies and assisted with
taking measurements. Dr. J. Robert Hunter allowed me to work at
Finca El Tigre and Dr. Ridgway Satterthwaite of the Associated Colleges
of the Midwest provided logistical assistance in various ways. Luis D.
Gomez of the Museo Nacional de Costa Rica provided a field vehicle.
Janice Mahlberg of the Milwaukee Public Museum provided photo-
graphic assistance. The assistance of Dr. Martyn Dibben and Neil
Luebke with growing peanut plants at the museum is greatly appreciated.
I also thank Joe Sugg, head of the North Carolina Peanut Council and
Dr. Harry Cobel (Crop Science, North Carolina State University) for
arranging peanut plants to be sent to me when the museum supply was
defoliated. I thank Cheryl Castelli for typing the manuscript. To all of
these people I am very grateful.

LITERATURE CITED

Baitey, L. H. 1969. Manual of cultivated plants. MacMillan Co., Toronto.

Levins, R. & R. MacArruur. 1969. An hypothesis to explain the incidence of
monophagy. Ecology 50: 910-911.

74 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Lewontin, R. C. 1965. Selection for colonizing ability. In The Genetics of Colo-

nizing Species (H. G. Baker and G. Stebbins, eds.). Academic Press, New York,
2 11-94.

Rien. L. D. 1968. The higher classification, phylogeny and zoogeography of the
Satyridae (Lepidoptera). Mem. Amer. Entomol. Soc., No. 24, 174 pp.

Orero, L. S. 1971. Instrucoes para criacao da borboleta “Capitao-do-mato”
(Morpho achillaena) e outras especies do genero Morpho (“Azul-seda’, “Boia’,
“Azulao-branco”, “Praia-grande”). Inst. Brasileiro Desenvolv. Florestal, Rio de
Janeiro, 27 p.

Tost, J. A., Jr. 1969. Mapa ecologico. Republica de Costa Rica. Centro Cientifico
Tropical, San Jose, Costa Rica.

Younc, A. M. 1972a. Adaptive strategies of feeding and predator-avoidance in
the larvae of the neotropical butterfly Morpho peleides limpida (Lepidoptera:
Morphidae). J. New York Entomol. Soc. 80: 66-82.

. 1972b. Community ecology of some tropical rain forest butterflies. Amer.

Midl. Nat. 87: 146—157.

. 1973. The comparative ethology and ecology of several species of Morpho

butterflies in Costa Rica. Studies Neotrop. Fauna 8: 17-50.

1974. The rearing of Morpho peleides (Morphinae) on peanuts. J. Lepid.
Soc. 28: 90-99.

Younc, A. M. & A. MuysHonpt. 1973. The biology of Morpho peleides in Central
America. Carib. J. Sci. 13: 1-49.

Journal of the Lepidopterists’ Society
32(2), 1978, 75-85

NOTES AND DESCRIPTIONS OF EUPTYCHIINI (LEPIDOPTERA:
SATYRIDAE) FROM THE MEXICAN REGION

Ler D. MILLER
Allyn Museum of Entomology, 3701 Bay Shore Road, Sarasota, Florida 33580

ABSTRACT. Several species of Euptychiini (Lepidoptera: Satyridae) are dis-
cussed and/or described. Described as new: Taygetis mermeria griseomarginata
(Guerrero, Mexico), Splendeuptychia kendalli (Tamaulipas, Mexico), and Cyllopsis
wellingi (Cayo dist., British Honduras). The previously unknown female of Cyllopsis
dospassosi L. Miller is described and figured.

Some years ago Mr. Roy O. Kendall of San Antonio, Texas sent me a
strange euptychiine satyrid from northern Mexico for identification. It
was apparent that the specimen was a representative of a new species in
the genus Splendeuptychia Forster (1964), a group hitherto known from
no further north than Panama. Since the Panamanian species, S. salvini
(Butler), was unrepresented in the Allyn Museum collection, I compared
the Mexican insect with the colored figures of salvini given by Butler
(1866) and by Godman and Salvin (1880 [1879-1901]). Many dis-
crepancies between the two insects became obvious, and a hurried call
to Mr. Gordon B. Small, Jr. of Balboa, Canal Zone resulted in his sending
two males of S. salvini that confirmed the superficial differences between
it and the Mexican butterfly as well as genitalic ones.

Once Mr. Kendall and his wife had managed to rear the Mexican
Splendeuptychia they needed a name on which to base the paper that
follows. Accordingly, I am taking this opportunity to describe the new
Splendeuptychia and some other euptychiines from the Mexican region.
Additional data are given on species that have come to my attention
since the publication of portions of my revision of the tribe. Some of
the new species described herein are members of genera not covered in
the revision yet, but it is felt that publication of these parts may be so
far in the future that other workers could benefit by having the names
proposed at this time.

Taygetis mermeria griseomarginata L. Miller, new subspecies
Figs. 1-5

Male: Head, thorax and abdomen clothed with dark brown dorsal and tan to
reddish-tan ventral hairs. Palpi reddish-tan, somewhat darker laterad. Antennae
dark brown dorsad, reddish-brown checkered with tan ventrad. Legs clothed with
brown hairs laterad, reddish-tan ones on inner portions of segments.

Wings with acutely falcate forewing apices as in T. mermeria excavata Butler
(1868). Upper surfaces of wings dark, rich brown, unmarked except for broad
(4-8 mm) grayish overscaling along margins of all wings and more narrowly and

76 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

1.0 mm

Figs. 1-5. Taygetis mermeria griseomarginata, n. ssp. 1-2, Holotype ¢ upper
(1) and under (2) surfaces; MEXICO: GUERRERO: Acahuizotla (Allyn Mus.
photos 101476-7/8); LFW (length of forewing) 47.0 mm. 3-4, Paratype 9 upper
(3) and under (4) surfaces; MEXICO: COLIMA: Comala (Allyn Mus. photos
101476-9/10); LFW 55.2 mm. 5, @ genitalia of Holotype; slide M-2732 (Lee D.
Miller).

VoLUME 32, NuMBER 2 Wale

less prominently along forewing costa. Under surfaces of all wings mottled in vari-
ous shades of brown, reddish-brown grayish-tan or ochreous (highly variable indi-
vidually) with forewing mesial bands poorly developed and only the extradiscal
bands of the hindwings well developed (usually delimited by some gray-green
scaling distad of the dark brown bands themselves); ocelli of both wings varying
from very well developed to obsolescent. Fringes of all wings gray above, tan to
reddish-brown below.

6 genitalia similar to those of other Mexican specimens (excavata) with some-
what stubbier valvae than those of South American representatives.

Length of forewing of Holotype ¢ 47.0 mm, those of the 21 ¢ Paratypes rang-
ing from 44.6 to 53.3 mm, averaging 50.35 mm.

Female: Similar in appearance to the ¢, differing chiefly in the paler coloration
both dorsally and ventrally and by the presence of a poorly defined transcellular
band of the hindwings beneath that is not shown by the ¢.

Lengths of the forewings of the seven 2 Paratypes range from 54.1 to 58.0 mm,
averaging 55.95 mm.

Described from 29 specimens, 22 males and seven females, from the western
slope of the Sierra Madre Occidental, Mexico.

Holotype ¢@: Mexico: Guerrero: Acahuizotla, ix.1957 (T. Escalante); @
genitalia slide M-2732 (Lee D. Miller).

Paratypes: all Mexico. GuERRERO: same locality as Holotype, 14 viii.1957, 5 4
ix.1957, 16 x.1957, 12 viii.1958, 12 iii.1958 (all T. Escalante); Tierra Colorado,
126 42 viii-ix.1971 (all A. Diaz Frances). Nayarir: vic. Compostela, 14
Posen bo Kiots). Comma: Colima, 14 11.1.1968; Comala, 14 31.x.1967,
12 14.41.1968 (all R. Wind).

Disposition of type-series: Holotype ¢, 17é¢ and seven @ Paratypes in Allyn
Museum of Entomology; single ¢ Paratypes will be placed in the American Museum
of Natural History, the National Museum of Natural History, Carnegie Museum
and the British Museum (Natural History ).

The name of this subspecies refers to the broadly gray dusted margins
of both wings on the upper surfaces. This situation is only hinted at in
specimens of T. m. excavata in which the maximum development of this
marginal gray scaling is about 1-1.5 mm on the forewing and virtually
absent on the hindwing. In the present subspecies this gray marginal
scaling is most prominent on the hindwing, but the forewing scaling is
more extensive than on any excavata specimen.

This gray-margined subspecies is apparently restricted to the western
slopes of the Sierra Madre Occidental from at least Nayarit to Guerrero.
While I have not seen material from all of the states in this area, I feel
confident that griseomarginata will be found in Jalisco, Michoacan and
possibly southernmost Sinaloa. A single specimen in the Allyn Museum
collection from Chiapas (Tuxtla Gutierrez, 13.viii.1961, leg. “M. S.”)
that was part of the Jae collection is referable to griseomarginata, but
since all material from Oaxaca and Chiapas that I have seen has been
referable to only excavata, I have excluded this Chiapas specimen from
the type-series. It may have been mislabelled, or it may represent a
genetic “throwback”, but it certainly is not typical of Chiapas-Oaxaca
before me. All of the specimens I have seen from the Nayarit to Guerrero

78 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

range have been referable to griseomarginata, and the presence of a single
specimen from outside this range should not be taken as “proof” that the
subspecies does not exist.

Splendeuptychia salvini (Butler), 1866
Figs. 6-8

Euptychia salvini Butler, 1866: 498 (type locality: Lion Hill, [Canal Zone],
“Panama” ).

Forster (1964: 128 ff.) erected the genus Splendeuptychia for 23 Neo-
tropical species, most of which are restricted to South America. The
species included in the present genus are among the loveliest of the
Euptychiini, and their pattern is unmistakable. Only S. salvini has thus
far been reported from Central America, and it is restricted to the Canal
Zone and adjacent Panama, possibly as far south as the Darien. S. salvini
seems to be a rare butterfly, at least in collections. I suspect that this
appearance of rarity is real, since the insect is one of the more spectacular
Satyridae and should not be overlooked by even a casual collector.

Two males were obtained from Gordon B. Small, Jr. for examination
and comparison with the Mexican species described below. The differ-
ences are cited under the new species, but it suffices to say here that the
two are not conspecific.

Splendeuptychia kendalli L. Miller, new species
Figs. 9-13

Male: Head, thorax and abdomen clothed with gray-brown dorsal and ochreous-
tan ventral hairs. Palpi pale gray laterad and dark gray ventrad and dorsad. An-
tennae brown dorsad, reddish-brown ventrad; tip of club slightly darker. Legs
clothed with gray hairs, but those of tarsi tan.

Wings above dull brown with three dark brown marginal lines separated by tan;
otherwise unmarked, but the markings of the under surface showing through
vaguely on this surface. Forewings below gray-brown in proximal half, tan in distal
half; two thickened rust-brown lines, one across cell, the other just outside cell
dividing the gray-brown from the tan ground color; three almost straight dark
brown marginal lines; between the marginal lines and the distal band is a row of
black-edged silver spots from M,—M:. to Cu:—2A, the whole spothand surrounded
by a thin brown ring. Hindwings below with gray-brown proximal and tan distal
ground color; thickened rust-brown bands of forewing continued on hindwing; three
thin, dark brown marginal lines following the slightly crenulate wing outline; a
mesial to submarginal yellow patch from Rs—M; to Cu.-2A encompassing silvered
spots in the interspaces, those in Rs—M:, M.-M2 and Cu,-Cu: with well defined
black irides; a subsidiary black line between the marginal lines and the yellow
patch from Cu, to the tornus; along M, and Cu; are two black submarginal patches.
Fringes pale gray above, tan below.

genitalia as figured, differing from those of S. salvini (Fig. 8) in many re-
spects, especially the shorter gnathos arms and the simpler valvae.

Length of forewing of Holotype ¢ 17.8 mm, those of the 374 Paratypes rang-
ing from 16.7 to 18.8 mm, averaging 17.67 mm.

VoLUME 32, NUMBER 2 79

0.5 mm

Figs. 6-8. Splendeuptychia salvini (Butler). 6-7, ¢ upper (6) and under
(7) surfaces; PANAMA: PANAMA: Bayano, nr. Pina (Allyn Mus. photos 010677—
11/12); LFW 15.9 mm; G. B. Small, Jr. collection. 8, 4 genitalia of same speci-
men; slide M-3418 (Lee D. Miller).

Female: Very similar to the ¢, differing chiefly in size, slightly paler coloration
and the more extensive yellow patches of the hindwing under surface.

Lengths of forewings of the 36 2 Paratypes range from 17.3 to 21.0 mm, averag-
ing 18.84 mm.

Described from 74 specimens, 38 males and 36 females, from the Mexican states
of Tamaulipas and San Luis Potosi.

Holotype ¢: Mexico: Tamauuipas: Gonzalez Ranch, nr. Los Kikos, ex ovum
on Bambusa aculeata, emerged 9.i.1975 (R. O. and C. A. Kendall); chromosome
specimen no. 3A-32-M; ¢ genitalia slide M-3651 (Lee D. Miller).

Paratypes: all Mexico. TamMau.ipas: same locality as Holotype, 2¢ 19
Momeni ovat LOV3., 194 M39 x 1973, 26 29 11974, Lea wl974, 36 49
xi.1974, 26 12 xii.1974, 16 792 i.1975, reared from Bambusa aculeata (all col-
lected or reared by R. O. and C. A. Kendall or W. W. McGuire). San Luis Porost:
Ciudad Valles, 29 vii.1970, 14 vii.1972, 36 39 vii.1973 (all collected by H. A.
Freeman); El Naranjo, 3¢ 1@ ii.1976 (all collected by R. O. Kendall); Tama-
zunchale, 1@ vii.1951 (T. Escalante).

10) JoURNAL OF THE LEPIDOPTERISTS SOCIETY

0.5 mm
ho

Figs. 9-13. Splendeuptychia kendalli, n. sp. 9-10, Holotype ¢ upper (9) and
under (10) surfaces; MEXICO: TAMAULIPAS: Gonzalez Ranch, nr. Los Kikos
(Allyn Mus. photos 010677-16/17); LFW 17.8 mm. 11-12, Paratype 2 upper |
(11) and under (12) surfaces; same locality as Holotype (Allyn Mus. photos

010677-14/15); LFW 19.6 mm. 13, ¢ genitalia of Holotype; slide M-3651 (Lee
D. Miller).

Disposition of type-series: Holotype 2, 10¢ and 109 Paratypes in the collec-
tion of the Allyn Museum of Entomology; nine ¢@ and 10 2 Paratypes returned
to R. O. Kendall; 194 and 1592 Paratypes returned to W. W. McGuire. These
series will be divided later among other museum collections.

I take great pleasure in naming this distinctive Mexican satyrid for

VoLUME 32, NUMBER 2 81

Mr. Roy O. Kendall who reared the Holotype and several other examples
in the type-series. His work on the life histories of various Mexican, as
well as Texan, butterflies has been of the greatest value to lepidop-
terology and promises even more future benefits to the science.

The Tamazunchale specimen came from the Escalante collection and
bore a cryptic determination label in an unknown hand identifying the
specimen as S. salvini. I had previously discounted salvini as the name
for the Mexican butterfly, and the receipt of true salvini confirmed my
previous analysis.

The genitalia are somewhat aberrant for members of Splendeuptychia
(Forster, 1964: figs. 161-164), especially in regard to the aborted
enathos.

In addition to the genitalic dissimilarities between kendalli and salvini,
the former may be distinguished by the following superficial characters:
1) the ground color of kendalli is browner, both dorsad and ventrad; 2)
the marginal lines on the upper surface are better developed in kendalli;
3) the ventral forewing of salvini bears four dark bands proximad of the
silver spotband, whereas in kendalli the basal of these is missing al-
together and the distal band is merely a thin line forming part of the
ring around the silvered spots; 4) the ventral hindwing of kendalli also
lacks the basalmost band that is prominent in salvini; 5) the yellow
patch of the ventral hindwing is more extensive in kendalli, whereas in
salvini this patch is poorly developed to absent posteriad of vein Cu,;
and 6) the silver spotband of the ventral forewing which extends
posteriad as far as 2A in kendalli reaches no further posteriad than Cu,
in salvini.

This species is apparently restricted to the mesic environments found in
a few places in the eastern foothills of the Sierra Madre Oriental. Thus far
the butterfly has been found in a very few localities from Tamazunchale
north to Tamaulipas where colonies of B. aculeata grow. What we know
about the bionomics of S. kendalli is given in a following paper (Kendall,
1978). Obviously, the insect is multivoltine.

Members of Splendeuptychia are almost uniformly rare. I suspect this
is a real occurrence, since they are much more attractive than are most
Euptychiini. Perhaps the relative abundance of S. kendalli and its as-
sociation with Bambusa will make possible the discovery of greater num-
bers of other species of Splendeuptychia. The association of this genus
with bamboo is further confirmed by S. S. Nicolay who brought me
several specimens of an as yet undetermined Splendeuptychia that he
took in a bamboo thicket in eastern Ecuador.

82 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Figs. 14-15. Cyllopsis dospassosi L. Miller, @ upper (14) and under (15)
surfaces; MEXICO: SAN LUIS POTOSI: El Salto Falls (Allyn Mus. photos
010677-2/3); LFW 17.7 mm; R. O. Kendall collection.

Cyllopsis dospassosi L. Miller, 1969
Figs. 14-15
Cyllopsis dospassosi L. Miller, 1969 (“1968”): 53; 1974: 84-86 (type locality:

52 mi. E Ciudad Victoria, Tamaulipas, Mexico ).

The type of this species remained unique until Mr. and Mrs. Kendall
collected one at El] Salto Falls, San Luis Potosi on 16.i.1975. This speci-
men is the second known example of dospassosi and, fortunately, is the
first female. It is quite comparable to the male, but the ground color of
the upper side is slightly darker, and that of the under surface is some-
what less olivaceous. Nevertheless, the maculation of the under surface
is comparable to that of the male with the addition of an ochreous outer
element to the extradiscal band of the hindwing. The “gray patch” en-
closing the ocelli of the ventral hindwing is obscure, as in the male. The
length of the forewing is 17.7 mm. I have not done a genitalic dissection
of this specimen since the female genitalia are not diagnostic in Cyllopsis
(L. Miller, 1974: 4).

The Kendalls’ specimen of this species (which is in their collection)
extends the known range of C. dospassosi from the Sierra de Tamaulipas
to the dry eastern flanks of the Sierra Madre Oriental, presumably of
Tamaulipas, as well as San Luis Potosi. The range of C. dospassosi may
be much wider than previously thought, and its rarity in collections may
be attributable to the usual lack of collecting of the smaller Euptychiini.

Cyllopsis wellingi L. Miller, new species
Figs. 16-20

Male: Superficially like C. nayarit (R. Chermock), but differing in the follow-
ing particulars: somewhat larger, approaching size of C. pephredo (Godman);

VOLUME 32, NUMBER 2 83

0.5 mm.
ne |

Figs. 16-20. Cyllopsis wellingi, n. sp. 16-17, Holotype ¢ upper (16) and
under (17) surfaces; BRITISH HONDURAS (BELIZE): Cayo District: Pine
Ridge, Thousand Foot Falls (Allyn Mus. photos 021777-1/2); LFW 17.3 mm.
18-19, Paratype 2 upper (18) and under (19) surfaces; same locality as Holotype;
(Allyn Mus. photos 021677-1/2); LFW 18.5 mm. 20, @ genitalia of Holotype;
slide M-3667 (Lee D. Miller).

84 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

under surface bands on both wings redder than in either species; transcellular bands
of both wings below less well developed than in nayarit; ochreous markings of hind-
wing below much more extensive than in nayarit (these are only hinted at in peph-
redo); and ocelli not edged inwardly with ochreous in gray patch area, as in nayarit.

4 genitalia as figured, not at all resembling those of pephredo. The genitalia do
bear some resemblance to those of C. pseudopephredo (R. Chermock) (L. Miller,
1974: fig. 141), a species that is otherwise quite distinct from wellingi and other
members of the pephredo subgroup in its lack of an androconial patch. The valvae
of the present species are somewhat broader than those of pseudopephredo, but the
characteristic inwardly directed teeth (which usually appear as dorsally diverted
ones) are very similar.

Length of forewing of Holotype ¢ 17.3 mm, those of the 11 ¢ Paratypes rang-
ing from 17.0 to 18.2 mm, averaging 17.48 mm.

Female: Differs from the @ of C. nayarit in much the same manner as does
the ¢, with the additional characteristic of a reddish flush on the upper surface
of some specimens that is not shown in other members of the pephredo subgroup.

Lengths of forewings of the eight 2 Paratypes range from 17.9 to 19.1 mm,
averaging 18.45 mm.

Described from 20 specimens, 12 males and eight females, from British Honduras
( Belize ).

Holotype ¢: British Honpvuras: Cayo District: Pine Ridge, Thousand Foot
Falls, 650 m, 2.ix.1976 (E. C. Welling M); @ genitalia slide M-3667 (Lee D.
Miller ).

Paratypes: all same locality as Holotype, 2—3.ix.1976 (E. C. Welling M.), 11é
82 (males all determined genitalically ).

Disposition of type-series: Entire type-series placed in Allyn Museum of Ento-
mology, but the series may be subdivided later.

It is with great pleasure that I name this little satyrid for Sr. Eduardo
C. Welling M. of Mérida, Yucatan, Mexico. He has consistently been
available to collect specimens of Euptychiini for me, and has often placed
undescribed and unexpected species at our disposal for systematic work.

This species is something of a puzzle. One male of thirteen in front of
me was a specimen of C. pephredo, but all of the others were the new
insect, as demonstrated by genitalic dissections. I have no idea why the
single pephredo was intermingled with wellingi at the type locality of
the latter, and this single specimen represents the first record of pephredo
from British Honduras. These two species now bring the total number
of Cyllopsis from that country to three (C. gemma freemani {Stallings
and Turner] also occurs there).

ACKNOWLEDGMENTS

I am most grateful to Messrs. Roy O. Kendall, E. C. Welling M.,
Gordon B. Small, Jr. and Alberto Diaz F. and Drs. W. W. McGuire and
Tarsicio Escalante for providing the material on which this is based.
Mr. A. C. Allyn took the photographs used to illustrate the paper. Mr.
Allyn and my wife and colleague, Jacqueline, read and suggested upon
the paper. To all of these individuals I owe a great debt of gratitude.

VoLUME 32, NUMBER 2 85

LITERATURE CITED

Butter, A. G. 1866. A monograph of the genus Euptychia, a numerous race of
butterflies belonging to the family Satyridae; with descriptions of sixty species
new to science, and notes on their affinities, &c. Proc. Zool. Soc. London,
[1866]: 458-504; ill.

1868. Catalogue of the diumal Lepidoptera of the family Satyridae
in the collection of the British Museum. London, Trustees British Mus.: vi +
211 pp.; ill.

Forster, W. 1964. Beitrage zur Kenntnis der Insecktenfauna Boliviens. XIX.
Lepidoptera III. Satyridae. Ver6dff. Zool. Staatssamml. Miinchen, 8: 51-188;
ill.

Gopman, F. D. & O. Satvin. 1879-1901. Biolcgia Centrali-Americana. Insecta.
Lepidoptera-Rhopalocera. London: 2 vols. + 1 vol. of ill.

KENDALL, R. O. 1978. Larval foodplant, life history notes and temporal distribu-
tion for Splendeuptychia kendalli (Satyridae) from Mexico. J. Lepid. Soc.,
32: 86-87.

Minter, L. D. 1969. On Mexican Satyridae, with description of a new species.
J. Res. Lepid., 7 (“1968”): 51-55; ill.

. 1974. Revision of the Euptychiini (Satyridae). 2. Cyllopsis R. Felder.

Bull. Allyn Mus., (20): 98 pp.; ill.

Journal of the Lepidopterists’ Society
32(2), 1978, 86—87

LARVAL FOODPLANT, LIFE HISTORY NOTES AND TEMPORAL
DISTRIBUTION FOR SPLENDEUPTYCHIA KENDALLI
(SATYRIDAE) FROM MEXICO!

Roy O. KENDALL?
Route 4, Box 104-EB, San Antonio, Texas 78228

ABSTRACT. Larval foodplant, Bambusa aculeata, Gramineae, rearing notes,
ecologic and temporal distribution at its northern distributional limit, are recorded
for Splendeuptychia kendalli Miller.

Field-collected adults of the satyrid, Splendeuptychia kendalli Miller,
were found at 2 locations: 1) along the Rio Sabinas at Rancho Pico de
Oro near Ciudad Mante, Tamaulipas, and 2) along the Rio Salto at El
Naranjo, San Luis Potosi. This species is closely associated with its
larval foodplant, Bambusa aculeata (Ruprecht) Hitchcock, Gramineae.
It is doubtless that this insect will be found at other locations along
water courses where its larval foodplant grows. Adults were taken in
January, February, July, October, November, and December over a 4-
year period. The areas were not visited during the other months. It is
therefore unknown whether this multivoltine species is continuous
brooded,; it may have a reproductive diapause.

Rearing. At Rancho Pico de Oro, 21 December 1972, I observed a 2
deposit a single egg on a juvenile leaf of B. aculeata. The ? was not
captured, but the egg was recovered and preserved.

Again at this location on 22 January 1974, two females were collected
and kept for egg production. Between 23 January and 2 February, 51
eggs were deposited in confinement on B. aculeata. Most of the eggs
were deposited by 1 female which died 2 February. The other female
was killed at this time, and both adults were preserved in alcohol to-
gether with 6 eggs. The remaining eggs hatched between 28 January
and 7 February. Larval losses were rather high resulting from an in-
adequate supply of fresh food. Attempts to keep the plants fresh in a
refrigerator were only moderately successful. Earlier, several specimens
of the foodplant were transplanted to the Los Arcos Courts gardens at
Ciudad Mante, our field headquarters, but they did not survive. In an
attempt to circumvent a 60-mile trip every few days for larval food, the
larvae were offered bermuda grass, Cynodon dactylon (L.) Pers. The

' Contribution No, 380, Bureau of Entomology, Division of Plant Industry, Florida Department
of Agriculture and Consumer Services, Gainesville 32602.
* Research Associate, Florida State Collection of Arthropods, Division of Plant Industry, Florida

Department of Agriculture and Consumer Services.

VoLUME 32, NUMBER 2 87

larvae ate the bermuda grass, and it was thought a laboratory solution
had been found for rearing this species. However, the larvae soon
began to die. The bermuda grass may have been toxic to the larvae,
but the lack of proper nourishment in the grass was suspect. On 6 April
1974 all remaining larvae (24) were preserved.

Once again a 2 collected 23 November 1974 from this location de-
posited 27 eggs between 24 and 29 November and died 3 December
1974. These eggs hatched between 28 November-l11 December, and a
maximum effort was made to rear them. Numerous trips were made to
the collection site for fresh bamboo. Even so, there were several larval
casualties attributed to rapid desiccation of the cut bamboo. Eleven
larvae pupated between 27 December 1974 and 12 January 1975. Adults
emerged (2 6, 72) between 7 and 23 January 1975. Two larvae and 2
pupae (one deformed ) were preserved.

Field-Collected Adults. In addition to the above, other field collections include:
Rancho Pico de Oro, 21 December 1972 (24,12), 9 January 1974 (32 ), 22 Janu-
ary 1974 (146), 22 February 1974 (14), 23 November 1974 (44, 19), 4
December 1974 (14), 6 December 1974 (39 ), and 8 January 1975 (1¢), all leg.
Roy O. and C. A. Kendall. At the same location, 27 December 1972 (124), 18 July
Poaee ee 20nfuly 1973 (2° ), 22 October 1973 (176, 82), 25 October 1973
(36, 22) all leg. W. W. McGuire. At E] Naranjo, 13 February 1976 (1@), 14
February 1976 (224), and 29 February 1976 (14) all leg. Roy O. and C. A.
Kendall.

ACKNOWLEDGMENTS

Mrs. Kendall and I wish to thank Sr. and Sra. Carlos Gonzales for
permission to conduct field research at their rancho, and for their warm
hospitality. To Sr. and Sra. Fernando Reyes Bugarin and their family
we are most grateful for the comfortable field headquarters provided
and for the use of their botanical gardens in our research.

Journal of the Lepidopterists’ Society
32(2), 1978, 88-96

NOTES ON THE LIFE CYCLE AND NATURAL HISTORY
OF VANESSA ANNABELLA (NYMPHALIDAE)

THomMas E. Dimock?
111 Stevens Circle, Ventura, California 93003

ABSTRACT. Observations on the life history of Vanessa annabella (Field)
show the early stages to be quite variable: the eggs in rib structure, and the later
larval stages in color pattern and behavior. Immature and adult behavioral charac-
teristics are similar to those of other Vanessa. V. annabella is usually present
throughout the year in coastal southern California.

Vanessa annabella (Field), the West Coast Lady, is a common and
familiar butterfly in western North America. Because it can usually be
found throughout the year in coastal southern California, opportunities
to study its life history are almost always present. However, there are
few published records available and none has included photographs of
the complete life cycle. Of published reports, Dyar (1889) gave one of
the more complete written accounts; Huguenin (1921) made some
general observations on the life cycle and natural history; and Coolidge
(1925) described the egg in detail and listed the larval foodplants. More
recently Emmel & Emmel (1973) illustrated paintings of a light form of
the last instar larva and the pupa and gave brief descriptive notes.

Specimens used for the present descriptions of the life cycle stages
were collected as freshly laid ova by following an ovipositing female at
the type locality in Ventura, California (Dimock, 1972). The leaves on
which these eggs were laid were placed in plastic containers 11 cm
square by 4 cm deep. Humidity was maintained by dampened tissue
paper placed on the container bottom. The containers were kept indoors
in a room temperature which varied from 17 to 25°C. Photographs and
measurements were made of each stage. Other specimens were reared
upon cut stalks of nettle placed in water so that leaf shelter construction
and other activities could be observed. Afternoon sunshine provided
direct and ambient light.

Full descriptions of the adults are given by Field (1971); thus, the
following adult descriptions are limited to those characteristics which
help distinguish V. annabella from related North American species.

Life Cycle Stages

_ Ege (lig. 1). Barrel-shaped, light green, with 10 to 14 transparent vertical
ribs. Measurements (Coolidge, 1925): 0.72 mm tall, 0.52 mm wide, tapering to
0.30 mm at base and 0.26 mm at top. Duration 4 days.

‘Museum Associate in Entomology, Natural History Museum of Los Angeles County, 900 Ex-
position Blyd., Los Angeles, California 90007. ;

VoLUME 32, NUMBER 2 89

Figs. 1-12. Vanessa annabella (Field): (1) egg, ca. 0.6 mm wide; (2) first
instar larva, 3 mm; (3) second instar larva, ca. 4 mm: (4) third instar larva, ca.
9 mm; (5) fourth instar larva, ca. 13 mm; fifth instar larvae, all ca. 30 mm: (6)
dark morph, black and yellow, (7) intermediate morph, orange and gray, (8) rusty
orange morph, (9) light morph, gray; (10) head, fifth instar, light tan morph;
(11) head, fifth instar, dark morph; (12) prepupa.

First instar larva (Fig. 2). Head shiny black, setae and thoracic legs black.
Ground color grayish brown after feeding for 2 days. When mature, body with
vague brownish mottling. Segments A-2, A-4, and A-6 with a pair of light yellow
spots between subdorsal and supralateral setae. Grows to 3 mm in 5 days.

Second instar larva (Fig. 3). Head shiny black. Ground color mottled dark
brown. Short branched spines black except for middorsal spines on A-4 and A-6
and subdorsal spines on A-2, A-4, and A-6, which are yellow. A narrow pair of
vague yellow lines divided by a narrow middorsal line of dark ground color
running from about T-1 to A-8. Grows to 4.5 mm in ca. 3 days.

Third instar larva (Fig. 4). Head shiny black with black setae arising from
black chalazae. Ground color usually black, but may begin to lighten as in lighter
morphs. Spines black except for subdorsal spines on A-2, A-4, A-6, and usually

90 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Wj

ee.
Z iY Wy,
» ty

Fig. 13. Vanessa annabella (Field): fifth instar larval nest on Urtica holosericea
Nutt.

middorsal spines on A-4 and A-6, which are yellow with black tips and yellow
bases confluent with dorsal double yellow lines. These paired lines, variable in
expression, separated by a middorsal line of ground color, are interrupted by
ground color at bases of non-yellow spines. Grows to ca. 9 mm in ca. 3 days.

Fourth instar larva (Fig. 5). Head black with bronze hightlights. Head cap-
sule width 1.5 mm. Ground color and markings nearly as variable as in fifth instar
(see following description). Grows to ca. 14 mm in ca. 3 days.

Fifth instar larva (Figs. 6-9). Head blackish or brownish black (Fig. 11)
with bronze highlights, or less often with vertical whitish tan stripes in light morphs
(Fig. 10). Capsule width 2 mm. Extremely variable in ground color and mark-
ings. Ground color varies from black to greenish white or grayish white, including
various browns and tans. Light markings present in fourth instar here vary from
dark rusty reds and oranges to yellow or various browns and tans. Extent of mark-
ings and lateral line varies independently of color; these tend to disappear alto-
gether in morphs of whitish ground color. Lateral line may be absent in any morph.
tusty orange spots may appear between subdorsal and supralateral spine bases,
varying in extent from absence to confluence with other pattern elements. Spines
branched, variable from black in dark morphs to whitish in light morphs, or dark
anteriorly and light posteriorly in intermediate morphs. Arrangement of spines:
middorsal row on segments A-1 to A-8, subdorsal rows on T-2 to A-8, supra-
lateral rows on T-2 to A-10, and lateral rows on A-1 to A-8. Body shape thickest
at midabdominal segments. Grows to ca. 25-30 mm in ca. 5 days.

Prepupa (Fig. 12). Light markings darken somewhat. Larva becomes slightly
shorter and thicker. Duration 1 day.

Pupa (Figs. 14-16). Head without projections. Mesothorax with a_ raised
middorsal point and a pair of subdorsal points. Metathorax with two large sub-

VOLUME 32, NUMBER 2 91

Figs. 14-20. Vanessa annabella (Field): (14) pupa, dorsal view, 19 mm; (15)
pupa, lateral view; (16) pupa, ventral view; (17) adult male, dorsal view, 47—mm
expanse; (18) adult male, ventral view; (19) adult female, dorsal view, 49-mm
expanse; (20) adult female, ventral view.

dorsal white spots, raised anteriorly. Abdomen with two small subdorsal white
spots on A-1 bordering those on metathorax. A-2 to A-7 each with one very small
middorsal point and a pair of more prominent subdorsal points. Color variable
from overall tan to mottled dark browns, sometimes with a greenish golden cast.

992 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

A spiracular line, darker brown than ground color, is variable in expression. Mea-
surements (average of five specimens): length 19 mm; width 7 mm; depths: thorax
6.25 mm, saddle 5.25 mm, abdomen 7.25 mm. Duration 8-11 days.

Adults (Figs. 17-20). Sexual dimorphism subtle, females having a more
rounded hindwing than males, especially at M:, Cu, and Cuz. Color pattern same
in both sexes: tawny orange with black markings, white subapical forewing spots,
and blue pupilled hindwing ocelli. On upperside, forewing cell crossed completely
by a black bar. Forewing costal bar, between cell end and apex, orange. Forewing
apex pointed, not rounded, at M,, with marginal tawny spot in interspace Rs.
Hindwing with four blue pupilled submarginal ocelli in interspaces M:, M2, Ms,
and Cu, and often a small solid black ocellus in interspace Rs. Hindwing under-
sides mottled principally in various buffs, tans, browns, and grays, with a whitish
triangle in interspace M» at cell end. Expanse averages between 40 and 48 mm,
females often larger than males.

Total developmental time for this species is ca. 30 to 36 days.

Natural History

Vanessa annabella uses a variety of foodplants in the families Urti-
caceae and Malvaceae. Native foodplants most frequently used in
southern California are Urtica holosericea Nutt. (Urticaceae), Sida spe-
cies, Sidalcea malvaeflora (DC.) Gray, and Sphaeralcea ambigua Gray
(Malvaceae). Introduced plants include Malva species, especially M.
parviflora L., and Althaea rosea (L.) Cav. (Malvaceae). John F. Em-
mel, M.D. (pers. comm.) also reports the use of Urtica urens L.
( Urticaceae ).

The eggs are laid singly, usually on the uppersides of the leaves. On
nettles (Urtica) the eggs are often attached to the sides of the stinging
spines.

The hatching larva eats away the top and adjacent walls of the egg
and crawls to a suitable place on the leaf uppersurface to construct a
shelter. This consists of fine silk webbing tied across a leaf midrib,
petiole, or small wrinkle on the leaf margin. The young larva lives under
this webbing as it feeds on the leaf and places its frass into the webbing,
creating a protective camouflage. In the second instar the larva may
enlarge the old nest or construct a new one nearby. When the larva is
at the growing tip of a nettle stalk, the nest may incorporate two or
more of the tiny new leaves. By the third instar the larva is capable of
folding a larger area of the leaf or constructing a deeper nest at the
petiole. In the fourth instar the entire leaf may be folded together (on
Urtica) or closed about the top edges (on Malwa). Frass is allowed to
fall out of the nest but often accumulates in piles in the nest bottom.
The fifth instar larval nest is usually larger and may incorporate neigh-
boring leaves and stems (Fig. 13). Sometimes leaves of nearby plants
which are not foodplants are also tied into the nest even though they are

VoLUME 32, NUMBER 2 93

not eaten. On plants with small leaves, such as young Malwa, the larva
may tie together many leaves before a nest enclosure is completed. On
Urtica holosericea, when a single leaf is used, larvae of V. annabella
usually construct nests on the uppersides of the leaves, either by folding
over one edge and securing it to the leaf surface or by tying both edges
together to form an enclosure. The petiole or nearby midribs may or may
not be partially cut to cause the leaf to hang vertically. Less frequently,
the larvae will fold the leaf edges underneath so that the undersurface
forms the nest interior.

Pupation sites are on either the foodplant or nearby objects. When the
foodplant is used, a leaf chamber is constructed with firm webbing and
the larva suspends itself from the chamber ceiling. Larvae in other loca-
tions may secure together any nearby objects to approximate an en-
closure or may simply pupate in exposed places, such as from twigs or
branches. Pupae often react to disturbances by wiggling laterally.

Emerging adults hang from the pupal shell or adjacent perch to ex-
pand their wings. A reddish brown meconium is ejected and the adult
is ready for flight in an hour.

Adults of V. annabella may be encoutered in any life zone from sea
level to alpine areas where open sunny places are preferred. Both sexes
visit flowers. In the afternoon males tend to congregate on hilltops or
other exposed places such as forest openings, glades, meadows, and
streamside slopes, especially when patches of dry, bare earth are avail-
able for sunning. Many man-made situations are particularly favorable:
windbreaks of trees, orchard rows, trails, firebreaks, garden paths, and
paved sidewalks and driveways. At these locations, when not occupied
in sunning, males will chase after each other and the other vanessid
butterflies Vanessa atalanta rubria (Fruhstorfer), V. cardui (L.), and
V. virginiensis (Drury), along with unrelated butterflies which con-
gregate in the same places. They often bravely chase larger insects and
birds and in general will investigate anything that flies through their
established area, including falling leaves and objects thrown overhead.
These activities ultimately bring the males into contact and subsequent
courtship and mating with females, but between these encounters the
males spend a great deal of time and energy simply chasing each other.
From observations made on hilltops in the vicinity of Ventura, California,
during November 1976 when all four Vanessa were present, it was noted
that any one species will chase the same or any other species, and two
or more individuals may join in the chase. The butterflies may chase
each other to a height of ca. 20 m or more before breaking chase and
quickly gliding down to land once again on the ground with backs to the

94 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

sun and wings spread. The butterflies’ wings frequently come into con-
tact during these encounters, but without damaging effects, and the re-
sulting noise can be heard nearby.

Females, although not congregating in the manner of the males, are
likely to be found anywhere, feeding, seeking foodplants, or ovipositing,
including hilltop localities when the foodplants or nectar sources occur
there also.

The flight of V. annabella is composed of glides with the wings held
horizontally, interrupted frequently by several fluttering beats. Chasing
is mostly vigorous fluttering, and the return dives are composed of
gliding and braking.

Diapause was not investigated, but if it does occur in this species it is
almost certainly during the adult stage, as it is in the other vanessid-
nymphalinid butterflies. If adults of V. annabella are unable to survive
prolonged or severe frosts, the species probably reinvades the greater
part of its northern and eastern range from the milder southwestern areas
where breeding is continuous throughout the year.

Vanessa annabella is easily attracted to suburban gardens by planting
Althaea rosea (Hollyhock) or encouraging Malva parviflora (Cheese-
weed) to become established. It is an easy butterfly to raise in captivity,
even under poor conditions. Larvae collected in the wild on one food-
plant (for example, Urtica holosericea) can be switched to other
foodplants (Malva, Althaea) when the former is less easily obtained.
The larvae are very often parasitized by tachinid flies, which emerge
from the mature butterfly larvae or pupae as mature maggots.

DiIscussIon

Coolidge (1925) noted that the egg ribs of V. annabella varied in
number from 11 to 13, with 11 the most common number. My ob-
servations confirm this, but eggs with 10 and 14 ribs were found during
the present study. This is in partial disagreement with Field (1971),
who stated that the genus Cynthia (in which annabella was placed) had
from 14 to 19 egg ribs, and Clench (in Howe, 1975), who gave 14 or 15
as the number of egg ribs in the subgenus Cynthia.

The larvae of V. annabella are extremely variable, as are the larvae
of V. atalanta rubria and V. cardui, and this variability not only makes
a written description difficult but compounds the task of providing re-
liable characteristics with which the three species can be separated. In
general, the descriptions given for the larvae in this article can be com-
pared with those for V. a. rubria and V. cardui in subsequent articles.
However, fifth instar larvae of V. annabella can always be distinguished

VoLUME 32, NUMBER 2 95

by their smaller head capsule width of 2 mm; in V. a. rubria and V.
cardui the fifth instar larval head capsule is nearly 3 mm. Once V. anna-
bella is identified, V. a. rubria is distinguished by the numerous white
cephalic chalazae, which in V. cardui are black.

From observations made on V. annabella larvae collected in various
locations and larvae reared under controlled conditions, it was noted
that the variations in ground color were at least partly due to environ-
mental conditions. The light, grayish-white morphs were more fre-
quently encountered on plants exposed to full sunshine, whereas the
darker morphs were found mostly on plants in secluded, shaded areas.
Darker morphs also resulted when larvae were reared under crowded
conditions.

Because Malva species are especially successful in disturbed areas and
are abundantly available throughout the year as foodplants, V. annabella
has probably become much more common since the introduction of these
weeds from Europe. This is the situation in the coastal southern
California lowlands, where a favorable climate prevails and V. annabella
can be found in every month of the year.

In his revision of the Vanessa butterflies, Field (1971) resurrected the
genus Cynthia for the carye, cardui, and virginiensis species groups.
Clench (in Howe, 1975) reunites all the species in Vanessa, treating
Cynthia as a subgenus. Emmel & Emmel (1973) used the cases of
hybridization between V. a. rubria and Cynthia annabella to demonstrate
the “close genetic relationship and probable generic identity” of the two
species. With no disrespect to the fine work of Field, I favor a treatment
similar to that of Clench, with reservations on the precise placement of
annabella. There are 10 known cases of hybridization between V. a.
rubria and V. annabella: one specimen reported by Edwards (1877),
one by Grinnell (1918), one by Gunder (1930), three by Dimock (1973),
one by Emmel & Emmel (1973), one specimen collected by Kirby in the
collection of the Natural History Museum of Los Angeles County, and
two specimens raised by Henne and Ingham in the Peabody Museum
collection. Mr. William D. Field (pers. comm.) discovered upon dis-
section the partially crippled specimen designated as “Hybrid #3” in
Dimock (1973) to be a female, not a male as erroneously reported. In
my opinion these occurrences support, at least, the arrangement of
Clench and the generic identity suggested by Emmel & Emmel. Biolog-
ically, the examples of hybridization may also demonstrate the presence
of an as yet incomplete reproductive isolatory mechanism caused by a
recent invasion of V. annabella from South America or a recent invasion
of V. a. rubria from Eurasia, or invasions by both species.

96 JoURNAL OF THE LEPIDOPTERISTS SOCIETY

ACKNOWLEDGMENTS

I wish to thank my father, M. W. Dimock, for printing the photographs
which accompany this article; Mr. Julian P. Donahue of the Natural
History Museum of Los Angeles County for access to the Museum col-
lection; Mr. Christopher Henne of Pearblossom, California, for an account
of his hybrids; Mr. William D. Field of the Smithsonian Institution,
Washington, D.C., for information on the hybrids; Dr. John F. Emmel,
Hemet, California, for reviewing the manuscript; and Mr. Alberto Muy-
shondt of El Salvador for recommendations on photographic techniques
and whose format for life history observations of El Salvador butterflies
is the one I followed for this article.

LITERATURE CITED

Cootmncr, K. R. 1925. California butterfly notes, III. Bull. Brooklyn Ent. Soc.
20(3): 146-147.

Dimock, T. E. 1972. Type locality and habitat—Cynthia annabella. J. Res. on
the Lepid. 10: 265-266.

1973. Three natural hybrids of Vanessa atalanta rubria x Cynthia
annabella (Nymphalidae). J. Lepid. Soc. 27: 274-278.

Dyar, H. G. 1889. Preparatory stages of Pyrameis carye Hiibner. Canadian Ent.
21: 237-238.

Epwarps, H. 1877. Pacific Coast Lepidoptera, No. 22. Notes on some diurnal
Lepidoptera, with descriptions of new varieties. Proc. Calif. Acad. of Scis. 7:
163-174.

EMMEL, T. C. & J. F. Emmeu. 1973. The butterflies of southern California. Nat.
Hist. Mus. of Los Angeles County, Sci. Ser. 26: 1-148.

Fretp, W. D. 1971. Butterflies of the genus Vanessa and of the resurrected gen-
era Bassaris and Cynthia (Lepidoptera: Nymphalidae). Smiths. Contribs. to
Zool., Number 84: 1—105.

GRINNELL, Jr., F. 1918. Some variations in the genus Vanessa (Pyrameis).
Psyche 25: 110-115, pl. 4.

Gunpver, J. D. 1930. Butterflies of Los Angeles County. Bull. Southern Calif.
Acad. of Sci. 29: 39-95.

Howe, W. H., coordinating editor. 1975. The Butterflies of North America.
Doubleday and Co., Inc., Garden City, L. I., New York. xiii + 633 p. + 97 pl.

HucuEnin, J. C. 1921. Life history of Pyrameis caryae in California (Lep.,
Rhop.). Ent. News 32: 216-217.

Journal of the Lepidopterists’ Society
32(2), 1978, 97-102

A NEW SPECIES OF HEMILEUCA FROM THE SOUTHWESTERN
UNITED STATES (SATURNIIDAE)

Pau M. TusKEs

Department of Environmental Toxicology, University of California, Davis.
Davis, California 95616

ABSTRACT. JHemileuca griffini Tuskes which occurs in southern Utah and
northern Arizona was collected for the first time in 1974. The adult moth is a black
and white day flying saturniid which is active during September and October. The
larval hostplant is black brush, Coleogyne ramosissima. This species has a unique
taxonomic position in that both the adult and larva exhibit morphological characters
which are intermediate to the Pseudohazis and Hemileuca groups, thus, a continuum
of characters exists between these two previously separated genera.

The genus Hemileuca consists of 23 described species, 16 of which
have partial or complete distributional patterns north of Mexico. The
moths within this genus are large to moderate in size, and exhibit a great
deal of hostplant and habitat diversity. Adults are characterized by hav-
ing the labial palpi fused to each other forming a small unsegmented
bilobed structure; also, the male has bipectinate antennae. Members of
Coloradia, the genus most closely related to Hemileuca, have labial palpi
which are separate, and males have antennae which are quadripectinate.

The last Hemileuca described as a distinct species was chinatiensis
(Tinkham), in 1943. The significance of H. chinatiensis as a species
with genitalic characters intermediate between Pseudohazis and Hemi-
leuca was overlooked by Tinkham; not until Ferguson (1971) was its
taxonomic position made clear. Michener (1962) combined the genera
Pseudohazis and Hemileuca on the basis of their morphological similarity,
but made no mention of chinatiensis. Although Michener included four
subgenera within Hemileuca Ferguson chose to abandon the subgeneric
names and to consider them as species groups.

It is the purpose of this paper to describe a new species of Hemileuca
collected for the first time in 1974, and to present additional mor-
phological evidence to support the merger of Hemileuca and Pseu-
dohazis. The new species of Hemileuca described in this paper is named
after Mr. Bruce Griffin, who collected the first specimens.

Hemileuca griffini Tuskes, new species

Holotype: Male (Figs. la,b). Heap: Eyes dark brown. Frontal and vertex
hairs rust red, clypeal hairs dark brown to rust red. Antennae, bipectinate, 0.67
cm long; shaft orange ventrally, dark brown dorsally, pectiniform processes black
and finely plumose. THorax: Dorsally clothed with black hairs; long white hairs
mixed with tufts of rust red hairs at base of secondaries. Collar, white with rust

98 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

red hairs anteriorly and posteriorly. Legs, clothed with black hairs; anterior por-
tion of pro- and metathoracic iegs with long rust red hairs. ABDoMEN: Abdominal
segments I-VI lustrous black, posterior margin of pleura I-VI lightly fringed with
white hairs. Terga I-VI black and sparsely clothed with red hairs. Terga VII and
VIII rust red. Forewincs: 2.54 cm long, wings approximately 60% white and
40% black. Veins black. Marginal area black, submarginal area white, but traversed
by black veins; postmedial line black and continuous. Distal portion of discal
cell with black band extending from postmedial line to costa. Coastal area black.
Area between costa and subcostal area adjacent to discal cell white. Antemedial
line black and extending from costa, curving out at base of discal cell and continu-
ing transversely to hind margin. Basal patch tear-shaped, black and free standing.
Basal portion below basal patch clothed with long black and white hairs to hind
margin of wing. Ventral surface similar to dorsal. Hrynpwines: 1.87 cm long
and approximately 50% white and 50% black. Marginal and submarginal areas
similar to those of primaries. Postmedial line diverging in area of the discal cell,
forming a circular area with a white center just distal to the discal cell. Basal area
black, clothed with long black and white hairs extending almost to the postmedial
line along the interior margin of wing. MALE Genirauia: (Fig. 5) Uncus trilobed,
dorsal and anterior portion covered with bristles; wide at base, narrowing at apex.
Lateral process of transtilla narrow, and not extending past apex of uncus. Transtilla
fused, but with shallow groove at apex of medial process. Valves prominent and
distinctly winged, apex slightly rounded and not pinnacle-shaped; upper half of
ventral margin heavily setose. Four—eight setae *4 as long or longer than juxta,
located basolaterally. Entire genitalic structure lightly sclerotized.

Allotype: Female (Figs. 2a,b). HEap: Eyes dark brown. Entire head covered
with rust red scales. Antennae, bipectinate, 0.70 cm long; shaft orange both dorsally
and ventrally pectiniform processes orange to dark brown, and not finely plumose.
THorAx: Dorsally clothed with black hairs, long white hairs at base of primaries
and secondaries. Long, large tufts of orange hairs at base of secondaries. Orange
hairs scattered on posterior portion of thorax. Collar rust red. Legs, similar to
those of male, but with more red present on femur of metathoracic leg than on
holotype. ABbpoMEN: Abdominal segments I—VII lustrous black with terga I—VII
lightly fringed with rust red hairs. Terga of segment VIII rust red. Pleura I—-VIII
lustrous black. Forewincs: 2.83 cm long. Similar to those of male, but with the
following exceptions: Marginal area more heavily marked with black scales. Black
margin continuous around entire wing. Hinpwincs: 2.04 cm long. Similar to
those of male but margins more heavily marked with black scales.

ParaTyPe VARIATION. The length of the forewing in the 20 males
examined averaged 2.54 cm, and ranged from 2.31 to 3.14 cm. The
markings on the forewings exhibited little variation, except for the black
basal patch. In some individuals the basal patch is slightly more
prominant than that of the specimen illustrated (Fig. 1), but in 15 of
the 21 paratypes examined it was less developed or almost absent. Al-
though the pattern is uniform, the intensity of the scales differ. The
wings of what are assumed to be older specimens are cream colored,
rather than white, and often partially transparent. The hindwings show
more variation than the forewings. Paratypes from Mexican Hat and
Bitter Springs Rd. appears similar, but two males from Pierce Ferry Rd.
are much darker. The forewings of 16 females examined averaged
2.75 cm, and ranged from 2.67 to 3.13 cm in length. As in the males the

VOLUME 32, NUMBER 2 99

forewings showed relatively little variation, but the hindwing differed
markedly. In most individuals the hindwings were approximately 50%
black and 50% white, while in others they were about 80% black.

Types: Howotyre: ¢ ca. 6 mi. S.W. of Mexican Hat, San Juan Co., Utah.
Elev. 4800’. Sept. 2, 1974. Bruce Griffin, Collector. ALLOTYPE: @ ca. 0.5 mi.
E. of Jct. 89A and 89 on Hwy 89, near Bitter Springs. Coconino Co., Arizona. Elev.
5200’. Collected as 3rd instar larva by B. Griffin and Ken Hansen on Coleogyne
ramosissima, May 3, 1975, and reared to maturity on Cercocarpa betuloides by
Paul Tuskes, emerged Oct. 12, 1976. Paratypres: Utah: 3 ¢ 2, same data as
holotype, 4 ¢@¢ and 3 2@Q same locality as type, Sept. 8, 1976, Kilian Roever;
8 646 and 4 22, Rt. 163, 7 mi. S.W. of Mexican Hat, San Juan Co., Sept. 8,
1974, K. Roever. Arizona: 3 ¢@¢ and 5 Q2@@Q same data as allotype; 2 ¢¢ and
3 29, Pierce Ferry Rd., 32 mi. N.E. Rt. 83, Mohave Co., K. Roever; 2 9 9,
Rt. 160, 42 mi. E.N.E. of Keyenta, Navajo Co., Sept. 8, 1974, K. Roever.

The types were deposited at the Los Angeles County Museum of Natural History.
Paratypes were deposited at the following institutions: American Museum of Nat-
ural History, Los Angeles County Museum of Natural History, Dept. of Entomology,
University of California, Davis, and the United States National Museum.

Larval Description—Last Instar

Head: Shiny black with numerous white setae, diameter, 4-5 mm. Clypeus
black. Body: Length 45-55 mm, width, 7-8 mm. Ventral surface gray to light
brown with an orange cast. Sublateral scoli black; lateral scoli black with slight
yellow cast at tips, dorsal scoli of rosette type, yellow with black center. Body
with three distinct lateral cream to white bands. Band I from prothoracic segment
to caudal segment, passing through sublateral scoli. Band II broken by interseg-
mental area and located slightly ventral to lateral scoli. Band III lightly pig-
mented broken band, but still obvious, located midway between dorsal and lateral
scoli. Segmental area with cream to white colored paniculum, especially common
on lateral areas. Secondary setae white. True legs black. Prolegs gray to black.
Spiracles orange. Ground color black.

Characteristics of H. griffini in relation to other Hemileuca

The trilobed uncus of griffini (Fig. 5) is typical of the Pseudohazis
group. The only Hemileuca to have a trilobed uncus outside of the
Pseudohazis group is H. electra Wright. The uncus of electra is typically
bilobed, but apparently some aberrant males have trilobed unci (Fer-
guson, 1971). The transtilla of griffini is shallowly grooved at the apex
of the medial process, and this characteristic appears to be intermediate
between the two subgenera. Fused transtilla are common to all Hemi-
leuca with the exception of diana and grotei. Both of these species have
bilobed transtilla which are rounded and short compared to the long
thin bilobed structure of Pseudohazis. The valves of griffini are prom-
inent and distinctly winged, showing much greater development than
the typical rounded valves of Pseudohazis but they are not as large as
those of most Hemileuca.

The genitalia and adult phenotype of griffini show the greatest

100 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Figs. 1-6. 1, Dorsal (la) and ventral (1b) view of male H. griffini (Holotype).
2, Dorsal (2a) and ventral (2b) view of female H. griffini (Allotype). 3, Dorsal
view of male H. chinatiensis.

t, Dorsal view of female H. chinatiensis. 5, Male
genitalia of H. griffini (Holotype). 6, Male genitalia of H. chinatiensis.

y)

VoLUME 32, NuMBER 2 101

similarity to those of chinatiensis (Figs. 3, 4 & 6). In griffini, the medial
process of the transtilla is frequently less sclerotized and not as stout
as that of chinatiensis. In addition, the medial process appears narrower
at the tip, with a slightly deeper groove than that found in chinatiensis.
The apices of the valves are variable: most are rounded, while others
have a more prominent constriction similar to, but less developed than,
those of chinatiensis.

Adult griffini are 25% smaller than those of chinatiensis. Phenotypi-
cally griffini males (Figs. la,b) can be distinguished from those of
chinatiensis (Fig. 3) in several ways: The basal black patch on the
forewing of griffini is usually free standing or almost absent, while in
chinatiensis the patch continues uninterrupted to the hind margin of the
wing. On the dorsal surface of chinatiensis the area between the costa
and radius adjacent to the discal cell is black while in griffini there is
a white patch between the costa and radius adjacent to the discal cell.
The viens proximal to the post medial line in griffini are narrow and
usually black; the veins of chinatiensis are also black, but the scaling
diffuses out from the veins giving them the appearance of being 3 to 4
times wider than those of griffini. The black margin of the forewings of
chinatiensis is from % to 1% times wider than that of griffini. In general,
the wings of griffini are approximately 40% black and 60% white while
in chinatiensis they are about 60% black and 40% white. The dorsal
portion of the abdomen of griffini may be covered with long rust red
and/or black hairs. Thus, the abdomen may vary from black to light
orange, with prominent rust red fringe around the anterior portion of
each tergum. The last two segments are covered with long red hairs
which form a tuft. The abdominal terga of chinatiensis are a uniform
rust red in color. These are but a few of the characters which may be
used to separate griffini males from those of chinatiensis.

Although the males of griffini and chinatiensis are very distinct, the
females are similar, with only a few obvious differences. The red
abdominal banding, prominent in chinatiensis females (Fig. 4), is less
developed or absent in griffini (Fig. 2). In griffini the red hairs are
mixed with black hairs and spread randomly over the entire terga but
become more abundant near the pleura, giving a diffused red appear-
ance to the lateral surface, or the abdomen may be completely black
except for the presence of short red hairs at the tip of the abdomen. At
present griffini and chinatiensis are though to be allopatric, with the
closest population of chinatiensis occurring 350 to 400 miles to the
southeast, in western Texas.

Examination of last instar griffini larvae indicated that they are similar

102 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

in most respects to Hemileuca. That is, secondary setae frequently, but
not always, arise from a pinaculum which is either white or cream
colored. Of 36 larvae examined, 31 had white pinacula, the remainder
were colored, giving them an appearance similar to that of Pseudohazis.
Thus, although most individuals appear similar to larvae of chinatiensis,
some individuals appear similar to H. hera (Harris). After examination
of the larvae of 16 of the 17 species of Hemileuca occurring north of
Mexico, this character has been found to be variable only in the larvae
of griffini.

The two genera, Hemileuca Walker, 1855 and Pseudohazis Grote &
Robinson, 1866 were joined by Michener in 1952. Michener based his
decision on external morphological characters. Ferguson (1971) showed
the intermediate characteristics of chinatiensis in which the genitalia are
similar to Hemileuca but the adult phenotype is that of Pseudohazis.
With the discovery and description of griffini, a second species with
intermediate adult and larval characters has been found. The genitalia
of griffini are more similar to genitalia of males of the Pseudohazis group
than those of chinatiensis. In addition, the larvae of griffini exhibit mor-
phological characters common to both species groups. Thus, a continuum
of adult and larval characters exists between the two previously separated
genera, Hemileuca and Pseudohazis.

ACKNOWLEDGMENTS

I would like to thank Mr. Bruce Griffin and Mr. Kenneth Hansen both
of Tucson, Arizona, and Mr. Kilian Roever and Mr. Michael Van Bus-

kirk of Phoenix, Arizona, for providing distributional data and/or speci-
mens.

LITERATURE CITED

Fercuson, D. C. 1971. The Moths of America North of Mexico. Fasc. 20.2A,
Bombycoidea (in part). Classey, London, pp. 101-153.

MicHeNER, C. D. 1952. The Saturniidae (Lepidoptera) of the Western Hemi-
sphere, Morphology, Phylogeny, and Classification. Bull. Amer. Mus. Nat. Hist.
98(5): 335-502.

TinkHaAM, E. R. 1943. Description and biological notes on a new saturniid of

the genus Pseudohazis from the Big Bend region of Texas. Can. Ent. 75(9):
159-162.

Journal of the Lepidopterists’ Society
32(2), 1978, 103-106

THE STATUS OF OLLIA PARVELLA DYAR: REDESCRIPTION
OF IT IN A NEW GENUS (PYRALIDAE)

ANDRE BLANCHARD
P.O. Box 20304, Houston, Texas 77025

ABSTRACT. Ollia parvella was described from females only. The discovery
of a few males shows that it is not a Peoriine but a Phycitine and that it belongs
in a new genus: Welderella.

When Shaffer (1968) revised the North American Anerastiinae (Auc-
torum), he grouped the majority of the species in the subfamily Peoriinae
and returned most of the remaining ones to the Phycitinae. A few genera
and species remained unplaced and were listed as such at the end of
his revision. Blanchard and Ferguson (1975) included three of these
unplaced species in the new phycitine genus Rostrolaetilia together with
seven new species which were described in the same paper.

Ollia parvella Dyar is another species which Shaffer could not place,
in this case because no male was available. My wife and I took six
specimens of this species (4 males and 2 females) 3 and 5 July 1975 at
the Welder Wildlife Foundation Refuge. Through the courtesy of Dr.
D. C. Ferguson I was able to borrow from the National Museum two
paratypes of the six females which constitute the type series. The
comparison of the genitalia of one of my females (slide A.B. 3879) with
those of one paratype (slide U.S.N.M. 52945) leaves no doubt that my
specimens are conspecific.

As Shaffer had suspected, an examination of the male genitalia shows
that this species is a Phycitine, although nothing closely related is in-
cluded in Heinrich’s revision of this subfamily (1956). Ollia is a synonym
of Peoria, not a Phycitine genus. Obviously a new genus is needed.

Welderella A. Blanchard, new genus

Type species: Ollia parvella Dyar (Figs. 1, 2, 4, 6).

Labial palpi porrect, downcurved, extending over three times eye diameter be-
yond front, loosely scaled; from beneath seen to be in contact with each other for
nearly all their length; second segment two and a half times longer than the third.
Maxillary palpi short, squamous. Antennae simple. No ocelli.

Forewing smooth, broadest at two thirds distance from base to apex; apex and
tornus rounded. Cell about two thirds length of wing. Venation somewhat vari-
able, 10 or 11 veins: R;, Rs and R; normally united, but two specimens show on
one wing R; separating from R;.; as a faint spur; M. and Ms; stalked for about two
fifths their length; Cu: from lower outer angle of cell; Cu. from near the angle.

Hind wing: length of cell ill defined (discocellular vein obsolete) but apparently
slightly longer than half the length of the wing; Sc and Rs long stalked, Sc separates
from Rs as short spur going io costa, Rs continues to near apex; M, straight to

104 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Figs. 1-5. Welderella parvella: 1, male; 2, female; 3, male genitalia; 4, de-
nuded labial and maxillary palpi, showing rudimentary tongue; 5, female genitalia.

outer margin; M. absent; M; and Cu, stalked for about half their length; Cu, in
almost exact prolongation of cubitus.

Male genitalia (Figs. 3, 7): Uncus triangular; apex produced, rounded, covered
with bristles. Gnathos stout, with very large flanged apical processes; the lobes
fusing posteriorly. Aedeagus smooth, moderately stout, about three times as long
as maximum width, vesica without cornuti. Tegumen with strong supporting struc-
ture forming on each side a wide U, with one branch supporting the gnathos and
the other the dorsal processes of the uncus. Vinculum mostly membranous, sup-
ported in part by the tegumen and the shallow, wide, well sclerotized saccus.
Transtilla incomplete, represented by a pair of irregularly shaped plates. Juxta
with anterior margin heavily sclerotized. Valves simple, without clasper; sacculus
narrow and short.

Female genitalia (Fig. 5): bursa and ductus bursae membranous; ductus bursae
rather wide, broadening progressively into pear shaped bursa; signum well sclero-
tized, longitudinally infolded; some weak scobinations around it; ductus seminalis
from left side of signum; genital opening wide, funnel shaped, sclerotized and
scobinate ventrally.

VOLUME 32, NUMBER 2 105

mam, 0.5 mm.

Figs. 6, 7. Welderella parvella: 6, venation; 7, enlarged part of male genitalia.

This genus shares characters with two widely separate groups of
Phycitine genera. The male genitalia suggest that it should go near
Laetilia: the uncus, the gnathos, the transtilla are quite similar, but the
complete absence of ocelli and the longitudinal wing pattern point to
a placement near Bandera and Tampa.

I take great pleasure in naming this new genus for the Welder Wild-
life Foundation, for its staff, whom I have always found ready to help
me in every way and for its generous founders, the late Rob Welder and
his wife Bessie Welder.

Welderella parvella (Dyar)

Dyar, 1906, p. 31. Barnes & McDunnough, 1917, p. 149. McDunnough, 1939, p. 36.
Kimball, 1965, p. 250. Shaffer, 1968, p. 89.

The original description reads: “Costal half of fore wing white with
slight darker lines on the veins toward apex. Inner half pale ocherous,
shading to gray next to white part. Hind wing whitish.” The Welder
Wildlife Refuge females match this description, but the hind wing of
the male is whitish only in the one third of it along the inner margin;
the other two thirds are blackish gray.

Wing expanse: males 12.5—-14 mm., females 13-15.5 mm.

Type data: I have not examined the holotype, a female from Brownsville, Texas,
June 3 (?), 1904, H.S. Barber, U.S.N.M. type No. 9103; genitalia slide No. 10,
Carl Heinrich, Dec. 20, 1932.

Specimens examined: Brownsville, Texas, 31 May 1904, 1 2; 8 June 1904,
1 2; (Slide U.S.N.M. No. 52945), both collected by H.S. Barber. Welder Wild-
life Foundation Refuge, 3 July 1975, 2 ¢¢4 (slides A.B. 3894, 3854, 3890; these
last two slides are all that remains of one of these males), 5 July 1975, 2 4 4,

106 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

2 292 (4 slide A.B. 3857, 2 slide A.B. 3879), all collected by A. & M. E.
Blanchard.

ACKNOWLEDGMENTS

I am indebted to Dr. D. C. Ferguson for arranging the loan of two
female paratypes from the National Museum collection and to Dr. J. C.
Shaffer for revising the manuscript.

LITERATURE CITED

Barnes, W. anv J. H. McDunnoucu. 1917. Checklist of the Lepidoptera of
Boreal America.

BLANCHARD, A. AND D. C. Fercuson. 1975. Rostrolaetilia—A new North Amer-
ican genus of the subfamily Phycitinae, with description of seven new species
(Pyralidae). Jour. Lepid. Soc. 29: 131-150.

Dyar, H. G. 1906. Description of new American moths. J. N. Y. Ent. Soc. 14:
30-31.

Hernricu, Cart. 1956. American moths of the subfamily Phycitinae. U.S. Nat.
Mus. Bull. 207, viii + 581 p.

Kimpati, C. P. 1965. Arthropods of Florida and neighboring land areas, I:
Lepidoptera of Florida, an annotated checklist.

McDunnoucu, J. 1939. Checklist of the Lepidoptera of Canada and the United
States of America, part 2 (Microlepidoptera). Mem. S. Calif. Acad. Sci., vol. 2,
no. l.

SHAFFER, J. C. 1968. A revision of the Peoriinae and Anerastiinae (Auctorum )
of America north of Mexico. U.S. Nat. Mus. Bull. 280, vi + 124 p.

Journal of the Lepidopterists’ Society
32(2), 1978, 107-110

BIONOMIC NOTES ON THE BLOOD-SPOT SKIPPER
[HESPERIIDAE: PHOCIDES LILEA SANGUINEA (SCUDDER) |

RAyMonp W. NECK

Pesquezo Museum of Natural History,
6803 Esther, Austin, Texas 78752

ABSTRACT. Observations on the life cycle of Phocides lilea sanguinea are re-
ported. An additional larval foodplant, notes on egg and larval stages, and adult
oviposition behavior are described.

Phocides lilea sanguinea (Scudder) is a large-sized skipper which
exhibits a metallic blue background with a prominent red spot on each
dorsal forewing. Basically a tropical species, sanguinea has established
breeding populations in the Brownsville, Cameron County, Texas area.
This population has been known as Phocides polybius lilea (Reakirt ) but
H. A. Freeman (in litt.) prefers treatment of lilea at the species level
with subspecific rank being accorded sanguinea. My initial interest in
this species arose because of the significance of establishment of per-
manent populations on non-native larval foodplants. Field observations
have yielded various new bionomic facts concerning this insect.

The only plant previously known to support larvae of sanguinea in
either the United States (Lipes, 1961) or Mexico (Comstock and Vasquez,
1961; Kendall and McGuire, 1975) is common guava, Psidium guajava
(Myrtaceae). My observations in Brownsville have revealed that a con-
generic plant, Psidium cattleianum Sabine (strawberry guava) is also
utilized as a larval foodplant by sanguinea. Identification of the food-
plant was verified using comparative herbarium specimens and _ char-
acters given by Bailey (1949: 729). All references below to Psidium refer
to P. cattleianum.

Newly-laid eggs are a pale but distinct aqua in color. A glistening
wet appearance is noticeable for several minutes following oviposition.
Within eighteen hours of oviposition, the contents of the egg begin to
turn reddish as embryogenesis proceeds. Red coloration appears initially
as individual foci which enlarge until the whole egg appears red. Similar
egg color changes during development have been reported in Agathymus
(Roever, 1964). This red color involves the internal constituents only, as
the chorion appears whitish in color. The egg is hemispherical in shape
with a diameter of approximately 1.5 mm. A hole encompassing the top
one-third of the egg reveals a hatched egg. The egg shell is not eaten
by the larva.

108 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Larvae of sanguinea go through a dramatic change in coloration during
development. First instar larvae have a bright red body similar in color
to the mature egg, but the head is brownish, varying extensively in
darkness. Body length is 3.0 to 3.25 mm while the head capsule measures
approximately 1.0 in width. Second instar larvae are about 8.0 mm long
with a head capsule of 1.5 mm in width. The head of the second instar
remains basically brown but is somewhat closer in coloration to the
red body color than is the head of the first instar larvae. Larvae of
Mexican and South American populations of Phocides were reported
to have yellowish intersegmental bands (respectively, Comstock and
Vasquez, 1961; Miles Moss, 1949); no such bands were observed on Texas
larvae. Larvae of intermediate instars were not observed. Mature larvae
are whitish with a slight “bloom” similar to that reported for Phocides
pygmalion okeechobee (Worthington) by Srohecker (1938). This
“bloom” is a white, powdery exudate which is present on the body of the
larva. The body also exhibits a large number of black “pin-prick” marks.
The head capsule is reddish-brown with a yellowish “eyespot” on each
side.

Retreats are formed by the larvae utilizing leaves of the foodplant.
Miles Moss (1949) described initial shelters as “small oval, of cutleaf”
while older larvae were “content to hide by day between several leaves
held together by a few strands of glutinous silk.” The following observa-
tions add to the above information: neither Lipes (1961) nor Kendall and
McGuire (1975) mention these retreats per se. Immature larvae make
two cuts from the leaf margin inward about five mm. This leaf section
is then pulled flat over toward the midrib so that a small retreat is
formed. This folded-over upper portion is then attached to the lower
portion by a plug of silk. The flatness of the retreat in comparison to
that formed by Calpodes ethlius (Stoll) on Canna is probably caused
by the thick stiff nature of mature Psidiwm leaves as opposed to the
thin, pliable leaves of Canna. While most retreats are formed from two-
leaves as reported by Comstock and Vasquez (1961), one retreat consisted
of three leaves—the apical pair and one of the leaves on the penultimate
node. The larva rested on the top leaf or “ceiling” in an upside-down
orientation.

Mortality is extremely high in the early instars. A check of a single
Psidium shrub yielded twenty-seven hatched eggs, but only five living
first and second instar (and dead bodies of seven other) larvae in larval
folds. Construction of the larval retreat is a task which many sanguinea
larvae are not able to complete. Leaves supporting hatched eggs but no

VoLUME 32, NUMBER 2 109

larvae occasionally have a single cut similar to the two required to form
a retreat.

Only a few chrysalids were observed during these studies. The chrysalis
is loosely attached to the upper part of the retreat by silk strands. A
parasitized chrysalis with many wasp (probably Apanteles) exit holes
was found in one retreat. This chrysalis measured 28 mm in length and
8.25 mm at greatest width (second abdominal segment ).

Adult female flight behavior in the vicinity of Psidium is quite dis-
tinctive. The imago “flits” or “skips” along and above the periphery of
the bush. Suddenly the adult will drop to the level of the bush and
quickly land on a leaf. Always facing outward toward the tip of the leaf,
she quickly lays an egg, and flies upward and around the bush again. If
undisturbed, the behavior sequence is repeated.

Although both male and female adults were observed flying, only
females alighted on any substrate. All but one of these observations in-
volved ovipositional landings on Psidium leaves. One female was ob-
served to land on a leaf of a bottlebrush shrub, Callistemon citrinus Staph.
(= lanceolatus DC.), which is also an introduced member of the Myrta-
ceae. After remaining on the leaf for about ten seconds, the adult flew
off; no egg was laid. The relatively long period of time spent on the
bottlebrush leaf indicates that after initial selection of a prospective
plant from a short distance (about 0.5 m) in the air, confirmation of
proper ovipositional substrate is required on the leaf surface. The means
of confirmation is unknown at this time, but it can be made in a very
short time if the substrate possesses the correct phytochemicals.

Placement of the eggs is highly predictable as a result of the rigid
ovipositional behavioral sequence. Of twenty-seven egg shells found
on 23 December 1970 all but one were on the upper surface of the leaf.
Terminal leaves tend to be selected for oviposition. Numerical data and
probable adaptive significance of this selection will be presented else-
where.

Adults in July were observed visiting flowers of an introduced orna-
mental, lilac-flowered golden dew drop (Verbenaceae: Duranta repens
L.). Adults expressed no interest in the flowers of the bottlebrush shrub
tentatively selected for oviposition.

Knowledge of the seasonal presence of the various life history stages
of sanguinea is desirable. Observations of a mature larva revealed no
apparent movement or feeding during a ten-day period from Dec. 1970-
Jan. 1971. Weather conditions at this time were quite mild; live eggs
and immature larvae were present at the same time. In contrast, De-
cember 1976 was cold and wet; no eggs or larvae were found on the
same bush at that time.

110 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

LITERATURE CITED

Bartey, L. H. 1949. Manual of cultivated plants. Rev. ed. Macmillan Co., New
York, 1116 pp.

Comstock, J. A. & L. G. Vasquez. 1961. Estudios do los circlos biologicos en
lepidopteros Mexicanos. An. Inst. Bil. Mex. 31: 349-448.

KENDALL, R. O. & W. W. McGuire. 1975. Larval foodplants for twenty-one species
of skippers (Lepidoptera: Hesperiidae) from Mexico. Bull. Allyn Mus. 27: 7 p.

Lives, J. E. 1961. More butterfly records from Brownsville, Texas, including a
foodplant of Phocides polybius (Hesp.). J. Lepid. Soc. 15: 114.

Mites Moss, A. 1949. Biological notes on some Hesperiidae. Acta Zool. Lilloana
7: 27-48.

Roever, K. 1964. Bionomics of Agathymus (Megathymidae). J. Res. Lepid. 3:
103-120.

STROHECKER, H. F. 1938. The larval and pupal stages of two tropical American
butterflies. Ohio J. Sci. 38: 294-295.

Journal of the Pepidonicnss Society
32(2), 1978, 111-115

CLIMATIC REGIMES RESULTING IN UNUSUAL
OCCURRENCES OF RHOPALOCERA IN
CENTRAL TEXAS IN 1968

RAYMOND W. NECK
Pesquezo Museum of Natural History, 6803 Esther, Austin, Texas 78752

ABSTRACT. During 1968 several species of Rhopalocera which normally do not
occur in this area (or occur only in small numbers during the latter part of the season )
were abundant in central Texas. Meteorological regimes which influenced this influx
of tropical species are discussed.

Central Texas is a major ecotonal area between the Nearctic and the
very northern fringes of a dilute Neotropical rhopaloceran element which
appears as far north as Central Texas under various climatic regimes.
Some of these species may occur as far north as Kansas or Nebraska as
stragglers, but breeding populations are not established. One such
climatic regime occurred in 1968. Discussed below are observations
made at the Brackenridge Field Laboratory of the University of Texas
at Austin within the corporate limits of the city of Austin.

Heliconius charitonius vasquezae Comstock and Brown and Dryas julia
moderata (Stichel) (both Heliconiidae) were very common in 1968 as
early as June. The existence of numerous fresh specimens and the length
of time during which these two species were present (well into the fall
months) indicate that breeding colonies of both species had become
established. At least one suitable larval foodplant, Passiflora lutea L.
( Passifloraceae ), is present on the grounds of the field laboratory. Noc-
turnal roosting aggregations (up to eleven individuals) of H. charitonius
were observed at several sites within the eighty-acre area. Different
sites were used at various times. Each site was a shrub or tree at the
edge of a wooded area. These two heliconians are seen in the Austin
area in about half the years, but they normally appear in late summer or
fall indicating late season dispersal from areas to the south with perma-
nent populations. Sporadic breeding occurs in at least some of these
years as indicated by reports for both species in 1966 (Rickard, 1967,
1968 ).

Dynamine dyonis (Geyer) occurred commonly from July into the fall.
Specimens were normally restricted to wooded areas along a dry arroyo.
Breeding occurred as indicated by the fresh condition of most specimens
seen throughout the season. D. dionis was initially reported in the Austin
area in 1899 (Brues, 1905). A breeding population was established
“along the bed of a dried up creek near Austin, Texas.” Brues had not

112 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

seen this species before this time (time of initial observations in Austin
area by Brues unknown) and did not see it during the three following
years, although he collected in the same areas. As his first records were
in October, this species probably immigrated into the area in late summer
1899; the progeny of these immigrants were observed by Brues. Brues
(1905) records that Mestra amymone (Menetries) was “very common
about Austin .. . in former years they were much less numerous.” M.
amymone is a permanent resident of the Austin area (Masters, 1970) but
was exceptionally abundant in 1968.

Adults of Achlyodes thraso tamenund (Edwards ) (Hesperiidae) were
commonly seen resting on soil surfaces in open areas in 1968. Kendall
(1965) reported this skipper from southern Texas. Records included
Goliad, Kleberg, Live Oak and San Patricio Counties (all well south of
the Austin area).

Hurricane Beulah struck the Texas coast near the mouth of the Rio
Grande River on 20 September 1967. Torrential rains covered a large
area of south Texas resulting in floods and semi-permanent ponds of
water (Grozier, et al., 1968; Baker, 1971). Subsequently, many rhopalo-
ceran species not before known from Texas (or the United States) were
reported from south Texas, particularly from Cameron and Hidalgo
Counties at the southern tip ( Doyle, 1970; Heitzman, 1970; Heitzman and
Heitzman, 1972; Kendall, 1970, 1972).

The effect of this storm upon the rhopaloceran fauna in south Texas is
well-documented and was more than temporary as some species have
been found in subsequent seasons (Tilden, 1974). While this storm may
have been related to the unusual rhopaloceran occurrences in central
Texas in 1968 in an indirect manner, the major cause involves the weather
of 1968. The species involved in this temporary northward movement
could have occurred in the central Texas area in late 1967 and remained
undetected, but their survival of the 1967-68 winter in these latitudes is
most unlikely. Coldest temperature at Austin for this winter was 22°F.
which is certainly much too cold for these species to survive. Average
monthly temperature for winter 1967-68 were generally below normal
in southern and central Texas. If these species could survive such cold
weather, these taxa would be common in central Texas during many
Se€aSons.

Torrential rains associated with Hurricane Beulah allowed the develop-
ment of larger than normal populations of these species in areas closer to
central Texas than is normally the case (southern Texas and/or northern
Mexico). The key to the appearance of these species in early 1968 was
higher than average rainfall in May 1968 at Austin (8.75” vs. normal

VoLUME 32, NUMBER 2 ans

4.22”) and many other localities in central Texas, especially along the
Balcones Escarpment (scattered localities actually had lower than normal
totals). Rains and cooler than normal temperatures during the summer
(U.S. Weather Bureau, 1968, Climatological Data, Texas, p. 73) fostered
plant growth and development of certain rhopaloceran populations.
Rainfall at Austin in summer 1968 was 18.5% above normal and high
temperature was only 98°F.

Population movements from south Texas or, more likely, from northern
Mexico along the Balcones Escarpment was initiated by large resident
populations; survival of these populations was allowed by favorable
moisture conditions. None of these forms was seen in central Texas in
early 1969 (low temperature during 1968-69 winter was 22°F.).

A second northward movement was observed in fall 1968. These move-
ments may have involved species more common in northern Mexico at
the end of the rainy season. Population development in northern Mexico
probably reached levels such that northward population movements
ensued. Climatic conditions of central Texas may not have been signifi-
cant in the occurrence of the following form as no evidence of repro-
duction in these relatively northern areas is known. Rainfall in the Border
Country of Texas was up to twice normal during this fall period (Posey,
1968; Wagner, 1969).

Colias (Zerene) cesonia (Stoll) is one of the common species in
Austin, particularly in spring and fall. This species is normally represented
in central Texas by the nominate subspecies which exhibits a reasonably
obvious “dog face” even in the female. Several female specimens of C.
cesonia collected during October 1968 exhibited a different phenotype,
immacusecunda Gunder, with greatly reduced black markings on DFW
and DHW. Originally described as a “form 2°” (Gunder, 1928), this form
has been treated as an aberration by some authors (Brown, 1965). I
believe this form represents a normal (seasonal?) phenotype present in
certain populations in Mexico (Neck, pers. obs.). Under certain climatic
conditions this form moves northward into Texas.

Other observers reported occurrences of tropical or sub-tropical species
in the central Texas area in 1968. Biblis hyperia aganisa Boisduval was
seen in San Antonio, Bexar County by W. Tyron on 7 October 1968
(Tilden, 1974), Kendall (1972) reviewed the occurrence of the tropical
heliconian, Eueides cleobaea zorcaon (Reakirt), in southern Texas as
far north as San Antonio in 1968.

In an attempt to verify that a particular climatic regime was responsible
for the occurrence of the above butterflies in central Texas, weather
records for 1899 and 1966 were consulted to determine if peculiar climatic

114 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

conditions existed at the time of previous occurrences in central Texas
of some of the above species.

In 1899, statewide precipitation in Texas was about normal, being
94% of average, but summer precipitation was high, 131% of average
(Norquest, 1941). Precipitation records for particular stations for 1899
are not published in readily available form. However, a massive rainstorm
occurred in late June 1899 in central Texas. This storm, which produced
the worst flood on record for the Brazos River, was centered north of
the Austin area, producing over thirty inches of rain in certain areas
(Texas State Almanac, A. H. Belo Corp.). If rain from this storm was
distributed, albeit in smaller amounts, to the south, then this storm with
its attendant rainfall and wind circulation could have been involved in
the occurrence of D. dyonis in central Texas in 1899. Ironically the oc-
currence of the subtropical butterfly D. dyonis in central Texas in autumn
1899 followed the most severe Texas winter on record. Record low
temperatures were recorded at various stations including 12°F. on 13
February at Brownsville at the southern tip of Texas. Central Texas tem-
peratures were at or below 0°F. Brues (1905) remarked that “the summer
had been very favorable for the development of insects as Hymenoptera
and Diptera were more abundant than I have ever seen them in that part
of the country.”

Occurrence of breeding H. c. vasquezae and D. j. moderata, in central
Texas in October 1966 (Rickard, 1967, 1968) probably resulted from
heavy rains in August of that year. Following below normal rainfall
during the first seven months of 1966, August was excessively wet for
both Austin (6.21” vs. normal 2.17”) and San Antonio (4.28” vs. normal
2.25’). Such a large influx of moisture is quite sufficient to cause rapid
vegetative growth and large scale movements of various butterfly pop-
ulations.

Environmental changes over large areas such as southern Texas and
northern Mexico provide natural experiments which reveal factors which
control the extent and abundance of occurrence of particular species (see
earlier report by Gilbert, 1969). Obviously, two major climatic regimes
of present-day central Texas prevent the establishment of a fringe Neo-
tropical fauna: insufficient rainfall and relative extreme winter cold.

LITERATURE CITED

Baker, ki. T., Jn. 1971. Relation of ponded floodwater from Hurricane Beulah to
ground water in Kleberg, Kennedy, and Willacy Counties, Texas. Texas Water
Development Board, 138: 33 pp.

Brown, K. S., Jn. 1965. Some comments on Arizona butterflies (Papilionoidea).
J. Lepid. Soc. 19: 107-115.

VoLUME 32, NUMBER 2 1S)

Burs, C. T. 1905. The occurrence of a tropical butterfly in the United States.
Entomol. News 16: 11-12.

Doyte, J. E., II. 1970. Field notes on three skippers in Texas (Hesperiidae). J.
Lepid. Soc. 24: 212.

GiutBerT, L. E. 1969. On the ecology of natural dispersal: Dione moneta in Texas
(Nymphalidae). J. Lepid. Soc. 23: 177-185.

Grozier, R. U., D. C. Haut, A. E. Hutme & E. E. ScHROEDER. 1968. Floods from
Hurricane Beulah in south Texas and northeastern Mexico, September—October
1967. Texas Water Development Board Report, 83: 197 pp.

Gunper, J. D. 1928. <A review of the genus Zerene Hbn. in the United States
(Lepid., Rhopalocera). Pan-Pacif. Entomol. 4: 97-102.

HeirzMan, J. R. 1970. A new U. S. butterfly record and a migration of Ewunica
monima in Texas. Mid-Cont. Lepid. Soc. 1(12): 10-11.

HerrzMan, J. R. & R. L. Herrzman. 1972. New butterfly records for the United
States (Hesperiidae and Libytheidae). J. Res. Lepid. 10: 284-286.

KENDALL, R. O. 1965. Larvae food plants and distribution notes for twenty-four
Texas Hesperiidae. J. Lepid. Soc. 19: 1-33.

1970. Three hairstreaks (Lycaenidae) new to Texas and the United States.

J. Lepid. Soc. 24: 59-61.

1972. Three butterfly species (Lycaenidae, Nymphalidae, and Heliconii-
dae) new to Texas and the United States. J. Lepid. Soc. 26: 49-56.

Masters, T. H. 1970. Distributional notes on the genus Mestra (Nymphalidae) in
North America. J. Lepid. Soc. 24: 203-208.

Norguest, C. E. 1941. Climate of Texas. In Climate and Man, Yearbook of Agric.,
pp. 1129-1146. U.S. Govt. Prtg. Ofc., Wash., D. C., 1248 pp.

Posey, J. W. 1968. The weather and circulation of September 1968. Monthly
Weather Rev. 96: 893-898.

Rickarp, M. A. 1967. An aberrant Heliconius charitonius (Nymphalidae). J. Lep.
Soc. 21: 248.

1968. Life history of Dryas julia delia (Heliconiidae). J. Lepid. Soc. 22:
75-76.

TitpENn, J. W. 1974. Unusual and interesting butterfly records from Texas. J.
Lepid. Soc. 28: 22-25.

Wacner, A. J. 1969. The weather and circulation for October 1968. Monthly
Weather Rev. 97: 88-94.

Journal of the Lepidopterists’ Society
32(2), 1978, 116-117

A NEW NAME FOR PAPILIO CERES CRAMER, [1776],
NEC FABRICIUS, 1775 (NYMPHALIDAE, DANAINAE)

GERARDO LAMAS

Museo de Historia Natural “Javier Prado,” Apartado 1109, Lima—100, Peru
and Department of Entomology, National Museum of Natural History,
Smithsonian Institution, Washington, D. C. 20560

ABSTRACT. Lycorea pieteri, nomen novum, is proposed as a replacement name of
Papilio ceres Cramer, [1776], pre-occupied by Papilio ceres Fabricius, 1775.

Papilio ceres, a danaine described by Cramer ([1776]: 141, 152, pl. 90,
fig. A) from “Surinam” is a junior primary homenym of Papilio ceres
Fabricius (1775: 504), an African nymphaline. The only author who
seems to have noticed this homonymy is Billberg (1820: 77). Under his
new genus Epimetes, Billberg introduced the name sebethis, as follows:

Sebethis Brasil. Eg. Ceres Fbr.

As can be surmised from other examples elsewhere in his work, the
above notation indicates that Billberg was proposing his name sebethis
as a substitute for ceres Fabricius, and that he had a specimen (or
specimens ) from “Brazil” in his collection (Eg. = Auctor hujus operis).
What is not clear is if Billberg considered ceres Cramer to be the senior
name, or if he intended to write “Cr.” instead of “Fbr.” after ceres, and
just made a lapsus calami. However, in at least one other instance,
Billberg gives preference to a junior Cramerian name over a Fabrician
one, under the genus Amaryssus Dalman (minos Cramer, [1780] versus
astenous Fabricius, 1775).

Whatever Billberg’s true intention was, it is his action which counts
here, and the result was the unfortunate introduction of an invalid junior
synonym of Papilio ceres Fabricius (currently known as Najas ceres
(Fox et al., 1965) or Euphaedra ceres Auctt.; the correct nomenclatorial
status of this species is not yet settled, cf. Cowan, 1974).

Therefore, Papilio ceres Cramer still needs a replacement name, and
under the provisions of Article 60(b) of the International Code of
Zoological Nomenclature I hereby propose pieteri, in the combination
Lycorea pieteri, nom. nov. This name is a masculine noun after the
patronym of Pieter Cramer.

ACKNOWLEDGMENTS

Mr. William D. Field, Smithsonian Institution, kindly read and com-

VOLUME 32, NUMBER 2 7.

mented on the manuscript. This paper was prepared during the tenure of
a post-doctoral fellowship at the National Museum of Natural History,
Smithsonian Institution.

LITERATURE CITED

Brtueerc, G. J. 1820. Enumeratio insectorum in Museo Billberg. [Holmiae], Gadel.
liv] + 138 p.

Cowan, C. F. 1974. Comments supporting the four outstanding requests affecting
butterfly generic names (Insecta, Lepidoptera). Bull. Zool. Nomencl. 30(3/4):
133-134.

Cramer, P. [1776]. De uitlandische Kapellen voorkomende in de drie Waereld-
Deelen Asia, Africa en America. Amsteldam, J. S. Baalde and J. Van Schoonhoven
& Comp.; Utrecht, Barthelemy Wild. 1(8): 133-155, pls. 85-96.

Fasricius, J. C. 1775. Systema Entomologiae, sistens insectorum classis, ordines,
genera, species, adiectis synonymis, locis, descriptionibus, observationibus. Flens-
burgi et Lipsiae, Korte. [iv] + [xii] + [xvi] + 832 p.

Fox, R. M., A. W. Linpsey, Jr., H. K. CLENcH & L. D. Mitter. 1965. The Butter-
flies of Liberia. Mem. Amer. Entomol. Soc. 19: [4] + ii + 438 pp., 1 pl., 233 figs.

Journal of the Lepidopterists’ Society
32(2), 1978, 118-122

TWO NEW PINE-FEEDING SPECIES OF COLEOTECHNITES
(GELECHIIDAE)

RONALD W. Hopces! AND ROBERT FE. STEVENS?

ABSTRACT. Two new species of moths, Coleotechnites ponderosae and C.
edulicola (Gelechiidae ), whose larvae mine needles of pinyon and ponderosa pine,
are described. Each is a potential pest species in Colorado and New Mexico.

Two species of Coleotechnites Chambers, one on ponderosa pine and
one on pinyon, are pests of these important tree species. Stevens has
studied both in the field, has published on the ponderosa pine species,
and has a paper on the pinyon species elsewhere in this issue. An attempt
at determination showed that each species is new. The senior author plans
a revision of the genus Coleotechnites, but this work will not be completed
for some years. Descriptions are presented to provide names for these
species.

Coleotechnites ponderosae Hodges & Stevens, n. sp.
(Figs. 1, 3, 4)

Upper surface as figured. Head: base of tongue mottled dark and medium gray;
labial palpus with first and second segments dark gray, nearly black on lateral surface,
second segment with a pale gray to white streak at 3% length and one at apex, mesal
surface white dorsally nearly to ventral margin, third segment white at base, middle
and apex, dark brown to nearly black separating white areas; frons mainly white, some
scales medium to dark-gray tipped; vertex and occiput with many dark-gray tipped
scales above eye, bases of scales white, many with lustrous purple reflections; scape
of antenna mottled white and very dark brown dorsally, white ventrally, shaft with
alternating half segments of yellowish-gray and brown scales on distal three-fifths.
Foreleg: mainly dark gray, some yellowish-white scales at apex of coxa, tibia with a
few white scales at half length and at apex; base and apex of first tarsal segment white,
apexes of second and third tarsal segments white. Midleg: similar to foreleg, some
white scales at base, middle and apex of tibia, base and apex of first tarsal segment
and apexes of second, third, and fourth tarsal segments white to off white. Hindleg:
coxa shining white and yellowish white with lustrous blue and purple reflections and
some gray scales; femur similar with gray scales on ventral margin; tibia with white
scales at base, a streak of white scales at base of first pair of spurs, and another at apex,
tibial spurs pale gray to off white, dorsal scale tuft yellowish white; tarsus with base
and apex of first segment, apexes of second, third, and fourth segments yellowish white,
other scales medium to dark gray brown. Wing length: 3.9-4.8 mm. Forewing:
upper surface mottled brown and white, scale bases white, fringe yellowish white in
tornal area. Hindwing: upper surface yellowish gray, fringe slightly more intense
yellowish gray; male with hair pencil of yellow to yellow-brown scales on posterior
margin at base. Ventral surface of wings mainly yellowish gray. Abdomen: dorsal

' Systematic Entomology Laboratory, IIBIII, Agr. Res. Serv., USDA, Beltsville, Maryland 20705.

e ; ane Forest and Range Experiment Station, Forest Sery., USDA, Fort Collins,
olorado )521.

VOLUME 32, NUMBER 2 119

2

a

Fig. 1, 2. Adult Coleotechnites. 1, C. ponderosae &, holotype; 2, C. edulicola ¢,
holotype.

surface shining yellow and yellowish gray with lustrous yellow and purple reflections,
male with yellowish-orange sex scales on segments 1-3; ventral surface mainly shining
gray, male with some white to off-white scales at apexes of segments 4-8, female
mainly darker gray. Male genitalia: as in Fig. 3; valvae asymmetrical, right valva
curved apically, left valva nearly straight, apex strongly curved; aedeagus ankylosed
with vinculum, apex reaching level of apexes of lobes of sicae; lobes of sicae directed
to left from middle to near apex, apex of right lobe turned posteriad; tegumen with
asymmetrical posterolateral lobes, right lobe slightly smaller than left lobe; gnathos
slender, hooked apically; uncus with posterior margin smoothly indented medially.
Female genitalia: as in Fig. 4; anterior margin of eighth segment heavily sclerotized;
base of ductus bursae slightly more heavily sclerotized than rest of ductus bursae:
ductus seminalis arising two-fifths length from base; corpus bursae gradually expanded
from ductus bursae: single signum with two, inwardly directed projections nearly
parallel sided for distal third, margins with small serrations.

120 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Foodplant: Pinus ponderosa Lawson. The larvae are needle miners. The life
history has been published by Stevens (1973).

Types: Holotype: ¢. Boulder, Colorado; July 1971; Hopkins No. US 36711;
Pinus ponderosa, J. Staley; USNM type number 75471. Paratypes: 6¢ 6, 829.
Same data as for holotype; USNM genitalia slides 10136, 37, 39-41, USNM.

Superficially, ponderosae is similar to the type specimen moreonella
(Heinrich); however, the wing length of moreonella is 6.2 mm, and the
basal segments of the antenna of moreonella are wider than long; those
of ponderosae are longer than wide. In the male genitalia the left valva
of ponderosae is very slender apically, and the apex is at a right angle
with the long axis; that of moreonella is stouter, and the distal third is
slightly curved. Coleotechnites ponderosae males have sex scales on
terga 1-3; moreonella males have them on segments 1-4.

Specimens in the type series vary in the number of dark-brown tipped
scales but generally all are dark brown as seen with the naked eye.
Worn specimens are paler brown.

Coleotechnites edulicola Hodges & Stevens, n. sp.
(mass 2, Sy, (6)

Upper surface as figured. Head: tongue mainly white with some gray scales at
base; labial palpus white at extreme base, lateral surface of first segment dark gray
beyond base to apex, mesal surface white, second segment mainly dark gray from base
to half length and with some dark gray-brown tipped scales at three-fourths length
laterally, a white band at half length and at apex, mesal surface mainly white with
some dark gray-brown scales on ventral margin, third segment mainly white with a
narrow brown band at half length; frons with dark-gray scales in front of eye and on
ventral half, dorsal part shining white to yellowish white; vertex and occiput mainly
white to yellowish white, many, scattered scales with gray apexes, with some lustrous
yellow and purple reflections; scape of antenna mainly white to yellowish white, some
gray-brown scales on dorsal surface before apex, shaft alternating half segments of
yellowish-gray and dark-gray scales. Foreleg: coxa mainly dark gray with some
yellowish-gray scales, femur darker gray brown with some white-tipped scales at half
length and on dorsal margin; tibia dark brown with white scales at one-fifth length,
one-half length and at apex; tarsus dark brown, base and apex of first segment and
apex of second segment with white scales, third and fourth segments with some
pale-gray scales at apexes. Midleg: coxa shining yellowish white with lustrous yellow
and purple reflections; femur mainly dark gray, some yellowish-gray scales at base
and one-half length and some yellowish-white scales at apex; tibia dark gray brown,
white at base, one-fifth length, three-fifths length and at apex; tarsus dark brown,
base and apex of first and apexes of rest of segments white. Hindleg: coxa shining
yellowish white and pale gray with lustrous yellow and purple reflections; femur white
on dorsal margin, dark gray with lustrous yellow and purple reflections on rest; tibia

with a broad white streak from base to nearly one-third length, a white streak at
three-fifths length, apex white, rest of segment dark gray brown, dorsal scale tuft
medium to dark gray, outer tibial spurs white, inner tibial spurs gray with white apexes;
tarsus mainly dark gray brown, base and apex of first segment and apexes of rest of
segments white. Wing length: 4.4-5.2 mm. Forewing: upper surface mottled dark

brown and white, fringe shining yellowish gray. Hindwing: gray, fringe shining
yellowish gray, a yellow to yellowish-brown hair pencil on posterior margin at base.

VoLUME 32, NUMBER 2 jal

Figs. 3-6. Genitalia of Coleotechnites spp. 3, C. ponderosae 6; 4, C. ponderosae
2:5, C. edulicola é; 6, C. edulicola.

Abdomen: ¢ with first three terga having yellow sex scales, other segments pale
yellowish gray, posterior margin of each segment paler, nearly white; ventral surface
mainly medium gray, posterior margin of most segments yellowish white to white;
2 much as for ¢ except lacking yellow sex scales. Male genitalia: as in Fig. 5;

122 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

right valva curved at right angle at one-half length and again at four-fifths length,
distal one-fifth in nearly same direction as basal half; left valva slender, tapering
gradually to acute apex; aedeagus extending slightly beyond apexes of sicae; tegumen
with pair of asymmetrical lobes on posterolateral margins; uncus broad, posterior
margin slightly indented medially. Female genitalia: as in Fig. 6; anterior margin
of eighth abdominal segment heavily sclerotized, anterior margin of eighth sternum
extending anteriorly to level of anterior margin of eighth tergum; ductus bursae
slender with a sclerotized band on basal one-fifth; ductus seminalis arising at one-
fourth to one-third length; corpus bursae expanded gradually from ductus bursae;
signum with pair of inwardly directed lobes; margins of each lobe very slightly serrate.

Foodplant: Pinus edulis Engelmann. The larvae are needle miners.

Types: Holotype: ¢. 14 Km N Aztec, New Mexico; 6/73; Pinus edulis; coll.
R. Stevens; Hopk. U.S. 36741; USNM type no. 75472. Paratypes: 12¢ 6, 32 9.
Same data as for holotype; USNM genitalia slides 3512-4, 10133-5, USNM.

Coleotechnites edulicola is very similar to ponderosae, but the forewings
are much paler. To the naked eye edulicola is a pale species; ponderosae
is a medium to dark gray-brown species. In edulicola the right valva has
two right angles, that of ponderosae is broadly curved and the apex is
at a right angle with the preceding part. Females of edulicola have the
anterior margin of the eighth abdominal sternum extending anteriorly
as far as that of the anterior margin of the eighth tergum; females of
ponderosae have the anterior margin of the eighth abdominal sternum
posteriad of the anterior margin of the eighth tergum.

Specimens in the type series vary somewhat in the relative amounts of
dark-brown and white scales. Coleotechnites edulicola has also been

reared from Pinus edulis at Santa Fe, and Nageezi, San Juan County,
New Mexico.

ACKNOWLEDGMENTS
We are indebted to Ann R. Richardson for the line drawings of the
genitalia and to Victor Kranz for the photographs of the adults.

LITERATURE CITED

STEVENS, R. E. 1973. A ponderosa pine needle miner in the Colorado Front Range. »
USDA For. Serv. Res. Note Rm-228: 1-3.

Journal of the Lepidopterists’ Society
32(2), 1978, 123-129

LIFE HISTORY AND HABITS OF COLEOTECHNITES
EDULICOLA (GELECHIIDAE) A PINYON NEEDLE
MINER IN THE SOUTHWEST

Rosert E. STEVENsS!, J. WAYNE BREWER”? AND DANIEL T. JENNINGS!

ABSTRACT. Coleotechnites edulicola infests needles of pinyon, Pinus edulis
Engelm., in the southwestern United States. The species is univoltine. Moths fly in
June and July, and eggs are laid inside previously mined needles. First-stage larvae
bore into green needles, feed within them, and overwinter there as 2d and 3d instars.
The insects pupate in late spring. Persistent infestations can cause severe defoliation
and presumably weakening and mortality of heavily infested trees. Several eulophids
and pteromalids, and a single species of braconid, are recorded as associates.

During summer 1973, our attention was called to a population of
needle miners causing heavy defoliation to pinyon (Pinus edulis Engelm. )
in the Animas Valley north of Aztec, San Juan County, New Mexico, and
on the Colorado side of the state line in La Plata County. We knew
that pinyon needle miners are occasionally reported from the Southwest;
the Aztec infestation offered an opportunity to make observations on the
life history and habits of the species.

Needle miners have received relatively little attention in western
North America. Freeman (1960) reviewed those of the entire continent,
discussing 23 species of gelechiids, yponomeutids, and tortricoids, mainly
from a systematic standpoint. The only ones that have been studied in
much detail in the West are two species of Coleotechnites (Gelechiidae )
infesting lodgepole pine, P. contorta Dougl.; C. milleri (Busck) in the
Sierra Nevada of California (Struble, 1972), and C. starki (Freeman)
in the Canadian Rockies (Stark, 1954, 1959). Two other species of
Coleotechnites, C. ponderosana Hodges and Stevens from Colorado
(Hodges and Stevens, 1978) and an undescribed species infesting Jeffrey
pine, P. jeffreyi Grev. and Balf. in southern California (Luck, 1976), have
also been studied, but in lesser detail.

DISTRIBUTION

We have reared C. edulicola from a number of northern New Mexico
localities, and have seen specimens reared from pinyon in southern Utah
(Fig. 1). We have also seen evidence of larval activity near Salida,
Chaffee County; Pueblo, Pueblo County; Walsenburg, Huerfano County,

1 Rocky Mt. For. and Range Exp. Stn., Fort Collins, Colo., 80521; the Station is maintained in
cooperation with Colorado State Univ. Jenning’s present address is USDA Bldg., Univ. of Maine,
Orono, Maine, 04473.

2 Dept. of Zoology and Entomology, Colorado State Univ., Fort Collins, Colo., 80523.

124 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

wwoeecbowa ee wwe o ooo
14 T
on | i]
iors 8
a !
fetes

"ema mk , '
$
,
j t
}
Peeemaeaeeeowaoae =
| 4
| i]
! t
!
'
t ‘
: 3
1
‘ ‘
Le

O

owns @
eT take

== 22 @2 22 @ Se @ @ aw ee

hae ee Tey

Fig. 1. Occurrence of Coleotechnites edulicola within general range of Pinus edulis;
@— adults reared, 0—larval feeding observed.

and Rio Blanco, Rio Blanco County, Colorado. The range of C. edulicola
may in fact coincide with that of pinyon over much of the Southwest. All
the rearings and feeding observations have been from naturally occurring

trees.

Lire History AND HABITS

Coleotechnites edulicola has a 1-year life cycle; the adults fly in early
summer and young larvae make up the overwintering stage (Fig. 2).

VOLUME 32, NUMBER 2 25)

Larvae

Larvae mm

x Larvaer -

Larvae

Feb. Mar. April May June July Aug. Sept. Oct. Nov. Dec. Jan.

Fig. 2. Generalized life history of Coleotechnites edulicola at Aztec, New Mexico.

The following details of the life history were developed from a series of
observations, collections, and rearings, mainly of the Aztec population,
during the period 1973-1976.

Adults

The adults are small silvery-gray moths, wingspan ca. 10 mm, found
from early June through mid-July. Hodges and Stevens (1978) present
a detailed description of both sexes. The moths are generally quiescent
during the daytime, rendered nearly invisible against twigs and bark by
their pattern of black and silver scales. When disturbed they fly rapidly
for a few seconds, generally within the branches of the tree on which
they were resting, and upon re-alighting, scurry rapidly to another resting
spot. Mating and oviposition have not been seen.

Eggs

The eggs are yellow-orange, nearly globular, and ca. 0.2 mm in
diameter. They are laid in clusters of variable size, around 6-12 eggs each.
Most eggs are laid inside needles mined out the previous year. A few
are also found in older mines. The eggs are generally within 1 or 2 mm
from an opening—either an exit hole or a “ventilation” hole.

126 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Teall Il IW

Number measured

| JI5 2 25.3 35 4 45 5 55 36mee
Head capsule widths (mm)

Fig. 3. Head capsule measurements (n = 894) of Coleotechnites edulicola larvae.

Larvae

The eggs begin hatching around the end of June (Fig. 2) and the
stage I larvae crawl to and colonize previously uninfested needles. The
larval period lasts most of the year, from early July until May or early
June of the following summer. Measurements of 894 head capsules
yielded a frequency curve with 4 distinct peaks, indicating 4 instars
(Fig. 3). Inspection of the curve might suggest stages I and II larvae
are members of the same set; however abandoned stage I head capsules
were readily found in mines containing stage II larvae, thus confirming
2 early larval stages with head capsule widths averaging ca. 0.15 and 0.25
mm, respectively. These observations on number of larval stages and
head capsule sizes closely parallel those reported by Luck (1976) on
the only other univoltine Coleotechnites studied in comparable detail.

VoLUME 32, NUMBER 2 7)

As shown in Fig. 2, C. edulicola was found to overwinter in both
instars II and III. Larva II is by far the most protracted stage.

Most (ratio ca. 5:1) larvae are found in needles of the current year’s
growth; i.e., most needles are attacked the year they are produced. A few
larvae infest older needles. Initial entry by stage I caterpillars is made
on the convex side of the needle, within ca. 2 mm of the apex. Early in
the larval period (e.g. 23 July 1975) mines are visible to the naked eye
only on close inspection. No more than a single larva per needle was
ever found.

As the larval period continues, additional needles of the current year’s
growth are invaded. Within the same needle bundle, larvae cross between
the flat (facing) needle sides; a small amount of silk often holds the
2 needles temporarily together. Larvae move to other needle bundles
without production of silk.

In the early larval period (instar I) frass is packed into the mined-out
needle, but in later stages it is pushed out. Needles inhabited by 3d or
Ath stage larvae have several holes for frass disposal and larval exit.

Throughout their lives the larvae are medium brown—most are
uniformly colored, others tend to be mottled. Head capsules and thoracic
and anal shields are dark brown to black. Fully developed larvae are
ca. § mm long.

Pupae

C. edulicola pupates around the end of May within the last mined
needle, near an open hole cut by the larva near the needle apex. The
pupae are elongate, cylindrical, black, and about 6 mm long.

EFFECT ON Host TREES

Repeated defoliation by Coleotechnites needle miners reduces growth
of stems, shoots, and needles, and—in more severe cases—kills trees (e.g.,
Struble, 1972). Such appears to be the general case with C. edulicola.
All degrees of damage, including tree mortality presumably resulting
from repeated defoliation, could be found at the Aztec site. Under the
circumstances (unmanaged southwestern pinyon-juniper forest), such
mortality does not constitute a problem. Only where pinyons are
intensively cultured and/or appearance is important would the species
likely be considered a pest.

Fig. 4 compares the appearance of shoots on a currently infested,
persistently defoliated tree with one that is essentially uninfested. The
infested needles are shed prematurely, leaving the shoots with a char-
acteristic bare-stemmed, tufted appearance.

128 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fig. 4. Branches from adjacent pinyons showing effects of severe (left) and
negligible (right) defoliation caused by Coleotechnites edulicola.

C. edulicola shares with C. ponderosana (Stevens, 1973) the character-
istic of colonizing trees in a highly differential manner. For example, the
shoots shown in Fig. 4 were taken at the same time from comparable
parts of 2 adjacent pinyons. One tree was severely defoliated (29.7%
of the needles infested) while infestation was negligible on the other.
Tree resistance may be involved.

ASSOCIATES

Several species of parasitic Hymenoptera were dissected or reared from
samples of pinyon foliage infested with C. edulicola from Aztec, Santa Fe,
and Nageezi (San Juan County), New Mexico. These included undeter-
mined species of Chrysocharis and Dicladocerus and a single specimen of
Zagrammosoma multilineatum (Ashmead), all Eulophidae, and several
pteromalids, not determined beyond family. A possibly undescribed
Apanteles (Braconidae) was reared abundantly from Aztec.

ACKNOWLEDGMENTS

We thank Les Eklund and Ron Shannon for field assistance. Associated
parasitic Hymenoptera were identified by Gordon Gordh (Eulophidae,
Pteromalidae ) and P. M. Marsh (Apanteles), U.S. National Museum,
Washington, D. C.

LITERATURE CITED

REEMAN, T. N. 1960. Needle-mining Lepidoptera of pine in North America. Can.
Entomol. Supp. 16, 51 p.

VoLUME 32, NUMBER 2 129

Hopces, R. W., & R. E. Stevens. 1978. Two new pine-feeding species of Coleo-
technites (Gelechiidae). J. Lepid. Soc. 32: 118-122.

Luck, R. F. 1976. Bionomics and parasites of a needle miner, Coleotechnites sp.,
infesting Jeffrey pine in Southern California. Envy. Entomol. 5(5): 937-942.
SrarK, R. W. 1954. Distribution and life history of the lodgepole needle miner
(Recurvaria sp.) (Lepidoptera: Gelechiidae) in Canadian Rocky Mountain

Parks. Can. Entomol. 86(1): 1-12.

1959. Population dynamics of the lodgepole needle miner, Recurvaria
starki Freeman, in the Canadian Rocky Mountain Parks. Can. J. Zool. 37:
917-943.

STEVENS, R. E. 1973. A ponderosa pine needle miner in the Colorado Front Range.
USDA For. Serv. Res. Note RM-228, 3 p. Rocky Mt. For. and Range Exp. Stn.,
Fort Collins, Colo.

StruBLE, G. R. 1972. Biology, ecology, and control of the lodgepole needle miner.
U.S. Dep. Agric. Tech. Bull. 1458, 38 p.

PIERIS NAPI OLERACEA (PIERIDAE) CAUGHT BY INSECTIVOROUS PLANT

Pieris napi oleracea Harris is frequently found in bog areas (Shull 1977, J. Lepid.
Soc. 31: 68-70) and swamps where insectivorous plants may occur. On 20 June
1977, Pamela Matthews, James Douglas, and I were collecting in a white cedar
(Thuja occidentalis L.) swamp north of Craftsbury, Orleans Co., Vermont. The
swamp contains sphagnum moss and small patches of sundew plants (Drosera
rotundifolia L.). In a sphagnum patch by our trail, we found a dead P. n. oleracea
female, which was caught by the dorsal surface of its body and wings on several
of the sticky sundew leaves. The external cuticle of the hapless butterfly appeared
to be intact, but the internal soft parts were gone. Because we had visited this area
twice during the previous week, and such a white object near our path would have
attracted our attention, we surmise that the butterfly had died recently. How it
became caught in the sundew, and whether its internal parts were digested by the
plant or by some sucking predator which encountered the immobilized butterfly
are not known.

Frances S. Cuew, Dept. of Biology, Tufts University, Medford, Massachusetts
02155.

Journal of the Lepidopterists’ Society
32(2), 1978, 130-134

MEYRICK’S RECORD OF “MECYNA FURNACALIS, GN.” FROM
FIJI, WITH A NEW GENERIC ASSIGNMENT FOR
PYRAUSTA HOMALOXANTHA MEYRICK
(PYRALIDAE: PYRAUSTINAE)

EUGENE MUNROE AND AKIRA MUTUURA

Biosystematics Research Institute, Agriculture Canada,
Ottawa, Ontario KIA OC6

ABSTRACT. Meyrick’s record of Mecyna furnacalis (Guenée) from Fiji belongs
not to Guenée’s species—described in Botys and now placed in Ostrinia—but to
Pyrausta homaloxantha Meyrick, described in 1933 from Fiji. This species is trans-
ferred to Xanthopsamma Munroe and Mutuura, a genus previously known from three
species from Japan, Korea, mainland China and Hainan. Figures of male and female
moths and genitalia are given and differential characters are noted. The apparent
isolation of the Fijian species may not be significant, as other species are likely to
turn up in the Indo-Papuan region.

Meyrick (1886: 264) identified a male and a female moth from Fiji
as Botys furnacalis Guenée, 1854: 332, and transferred the species to
“Mecyna, Gn.,” saying, “This species agrees well enough with Guenée’s
description, but as that is in some respects incomplete, I have redescribed
it to prevent error. ... Guenée’s type is stated to be from Australia, but
I think this is probably an error, and may be neglected until confirmed.”

Mr. Michael Shaffer has called our attention to a male specimen from
Fiji in the British Museum (Natural History), which for many years
was labelled “Type” over the name label “furnacalis Meyr.” This is
evidently the male recorded by Meyrick (1886), as we have been unable
to trace any furnacalis named by Meyrick. Meyrick’s identification,
which we overlooked when we referred Botys furnacalis Guenée to
Ostrinia in our revision of that genus (Mutuura and Munroe, 1970: 33),
is erroneous. We consider the specimen conspecific with Pyrausta
homaloxantha Meyrick (1933: 411), described from a female holotype
from Vunidawa, Fiji, in the British Museum (Natural History). The
maculation is virtually identical in the two specimens (Figs. 1, 2), the
localities agree sufficiently, and as will be shown, the genital characters
of the two sexes are concordant.

The species is not properly referable to Pyrausta Schrank, 1802,
(= Botys Latreille, 1802-1803), to Mecyna Doubleday, 1850, to Mecyna
in the sense of Guenée, 1854, (= Uresiphita Hiibner, [1825]) or to
Ostrinia Hiibner, [1825], all of which differs more or less widely in
genital characters (see Munroe, 1950, 1976). Instead, the species is

VoLUME 32, NUMBER 2 131

Figs. 1, 2. Xanthopsamma homaloxantha (Meyrick), specimens in British Museum
(Natural History). 1, ¢, so-called type of “furnacalis Meyr.”; 2, 2, holotype of
Pyrausta homaloxantha Meyrick.

132 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fig. 3. Xanthopsamma homaloxantha (Meyrick), ¢ genitalia of specimen illu-
strated in Fig. 1.

clearly a member of the genus Xanthopsamma Munroe and Mutuura,
1968, previously known from a complex of three species from Japan,
Korea, mainland China and Hainan. The Fijian species agrees in external
structural characters and in the essentials of male and female genitalia
with the type-species, X. aurantialis Munroe and Mutuura, 1968, and its
previously recognized relatives.

In maculation X. homaloxantha (Meyrick), n. comb., differs from its
temperate east Asian congeners in its yellower forewing, without terminal
infuscation, with finer but more distinct transverse lines, and with
relatively weaker discocellular bar, and in the contrastingly whitish-
yellow, not almost concolorous buff or fulvous hindwing, without post-
medial line or terminal infuscation. The termen of the forewing is less
strongly curved than in the previously included species and the wing
consequently appears wider and its apex sharper.

The male genitalia (Fig. 3) are closely similar to those of X. aurantialis
and X. youboialis, but have the uncus relatively narrower and sharper,
the dorsal prominence of the sacculus longer and more gradually rounded,
and the spinulose zone at the base of the basally directed lobe of the
clasper somewhat wider. The female genitalia (Fig. 4) resemble those
of X. aurantialis, but have the tapering part of the ostial chamber shorter
and separated from the collar of the ductus bursae by a narrow un-
sclerotized zone, and have the signum relatively larger.

At present no great significance should be attached to the apparent
wide disjunction in the geographical range of the genus, as further species
may well exist in tropical Asia and Melanesia.

VOLUME 32, NUMBER 2

133

\\

Mi NN
wi my .

ir:

o
bom.

rere i

4

Fig. 4. Xanthopsamma homaloxantha (Meyrick), 2 genitalia of holotype
Fis 2).

(see

134 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

ACKNOWLEDGMENTS

We thank Mr. Michael Shaffer and the authorities of the British
Museum (Natural History) for information and the opportunity to study
the material discussed. The photograph of male genitalia was made by
Mr. T. Stovell, Graphics Unit, Agriculture Canada, with the assistance of
Mr. D. H. Kritsch. The drawing of female genitalia was made by Mr.
Arthur Smith.

LITERATURE CITED

GuENEE, A. 1854. Species général des Lépidoptéres. 8. Deltoides et Pyralites.
Paris.

Meyrick, E. 1886. Descriptions of Lepidoptera from the South Pacific. Trans.
Entomol. Soc. Lond., 1886, pp. 189-296.

1933. Pyraustidae. Exot. Microlepid. 4: 393-411.

Munroe, E. 1950. The generic positions of some North American species com-
monly referred to Pyrausta Schrank (Lepidoptera: Pyralidae). Can. Entomol.
82: 217-231.

. 1976. In Dominick, R. B., et al. The moths of America north of Mexico,

Fasc. 13.2. London.

, & A. Muruura. 1968. Contributions to a study of the Lepidoptera of
temperate east Asia. III. Can. Entomol. 100: 974—985.

Muruura, A., & E. Munroe. 1970. Taxonomy and distribution of the European
corn borer and allied species: genus Ostrinia (Lepidoptera: Pyralidae). Mem.
Entomol. Soc. Canada, 71.

SCREECH OWL PREYS ON PERIDOMA PLECTA (NOCTUIDAE)

As part of a long term study of populations and mortality of the Screech Owl
(Otus asio) I collected a vehicle killed specimen near Oxford, Connecticut on 4 May
1976. Routine measurements were taken and stomach contents analyzed. Eight
larvae of the Flame-shouldered Dart, Peridroma plecta L. (Noctuidae) were found
among the owl's stomach contents. The P. plecta larvae were fresh and readily
identifiable, suggesting that the owl was killed shortly after feeding, but before
digestion had begun. Although Bent (1938, U.S. Nat. Hist. Mus. Bull. 168. 482 p.)
and others have recorded a variety of Lepidopteran adults and larvae as occasional
Screech Owl prey, this constitutes the first record of such for noctuid species. This
observation indicates the susceptibility of noctuid larvae to efficient nocturnal
predators. It also provides absolute evidence that at least one species of large, avian
raptor will feed opportunistically on available insect larvae.

Dwicur G. Smrru, Biology Department, Southern Connecticut State College, New
Haven, Connecticut 06515.

Journal of the Lepidopterists’ Society
32(2), 1978, 135-137

“ORANGE” BANDS, A SIMPLE RECESSIVE IN
ANARTIA FATIMA (NYMPHALIDAE)

ANNETTE AIELLO! AND ROBERT E. SILBERGLIED!

Smithsonian Tropical Research Institute,
P.O. Box 2072, Balboa, Panama Canal Zone

ABSTRACT. Breeding experiments with aberrant Anartia fatima butterflies having
Orange instead of red hindwing markings produced results consistent with the inter-
pretation that the orange color is due to the homozygous condition of a simple
Mendelian recessive allele.

We report here on the genetics of a color aberration in the common
Central American nymphalid butterfly Anartia fatima Fab. Throughout
its range, from southern Texas south into eastern Panama, A. fatima bears
on the hindwings a conspicuous, elongate red band, approximately 9 mm
long by 2 mm wide (Fig. 1, arrow). Wing pattern is identical in males
and females, and it is usually necessary to examine the genitalia in order
to distinguish the sexes.

Butterflies in which the hindwing markings are orange instead of
red (Fig. 1, right) appeared among the progeny of a phenotypically “red”
female from Achiote, Colon Province, Panama. This individual had been
brought to Barro Colorado Island in the Canal Zone during June 1977 as
part of a larger study involving the rearing of many A. fatima for behavior
and genetic research. Among her 91 offspring, 21 had orange bands,
a ratio of 3.33:1, “red”:“orange” butterflies.

Using these offspring, we obtained the crosses shown in Table 1.
“Orange” bred true. Two “red” to “red” crosses produced entirely “red”
individuals, while one other “red” to “red” cross produced a ratio of 3.53:1
“red’:“orange’ individuals. Three “orange” to “red” crosses produced
44 “red” butterflies, with no “orange” appearing.

These results are consistent with the interpretation that a simple
Mendelian recessive allele (r) is involved, with a double dose (rr)
resulting in the “orange” phenotype. The “red” female from Achiote
must have been heterozygous for “orange” (Rr) and so must have been
her mate. The same would be true of the two “red” pairs which pro-
duced the 15 “orange” out of 68 offspring.

A. M. Shapiro (pers. comm.) has seen an orange variant in A. amathea
at Cali, Colombia. Considering both the partial genetic intercom-
patibility of A. fatima and A. amathea (Silberglied and Aiello, in prep. }
and the intermediate expression of red pigmentation among hybrids,

1 Present address: Department of Biology, Harvard University, Cambridge, Massachusetts 02138
S.A.

136 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

no filter

green filter

Fig. 1. “Red” (wild type, left) and “orange” (aberration, right) phenotypes of
A. fatima (males). The arrow (upper left) indicates the characteristic colored band.
The lower photographs were taken through a green filter (Tiffen green no. 1) to
enhance the contrast between the bands of the two phenotypes.

VoLUME 32, NUMBER 2 137

TaBLE 1. Crosses involving the “orange” phenotype of Anartia fatima.

Phenotypes! Phenotypes
of parents of progeny Papa’ ratio,

——_—_—— Number of — R:O, if a simple
Q of crosses “red” “orange” recessive p?
R® I 1 70 21 381) 30)
R° R° 1 53 15 Osh So
R R 2, 38 0 1:0 -
OF R 3 44 0 1:0 -
O° (ig J) 0 78 0:1 _

1R = “red,” O = “‘orange.”’

2 Probability (Chi-square test, 1 degree of freedom) that differences between expected and
observed values are due to chance alone.

3 Female collected in Achiote, Panama.

4 Phenotype of male unknown; presumed (based upon results) to be “‘red”’ (and genotype pre-
sumed to be Rr).

> Progeny of female collected in Achiote, Panama.

that variant is likely to be genetically homologous with the one reported
here.

Specimens of this aberration have been deposited in the collections of
the Museum of Comparative Zoology, Harvard University; Peabody
Museum of Natural History, Yale University; American Museum of
Natural History, New York; National Museum of Natural History, Wash-
ington, D.C.; and G. B. Small, Panama Canal Zone.

Journal of the Lepidopterists’ Society
32(2), 1978, 138-139

A GYNANDROMORPH OF PAPILIO POLYXENES (PAPILIONIDAE )

Cultures of Papilio polyxenes asterius Stoll are maintained in our laboratory for use
in ecological studies. In October 1975, a bilateral gynandromorph of this species ap-
peared in the second generation of a laboratory culture derived from populations
around Brooktondale, Tompkins County, New York. This was the first such specimen
observed.

Although the external genitalia are male, only the right half of the specimen is
male in appearance (Fig. 1). The yellow spots of the inner row of the right forewing

Fig. 1. Lab-reared gynandromorph of Papilio polyxenes, dorsal view.

are greatly reduced and/or appear as “ghosts” composed of scales intermediate in
color between black and yellow. The inner band of the hind wing resembles a normal
male except for the last yellow spot (Cu: cell) which is half obliterated by black scales.
The hazy blue spots of the hind wing are irregular and appear in cells Cui, Cue, Me,
and Ms. The left half of the specimen perfectly resembles a female. The left forewing
is 43 mm in length, the right is 41 mm.

Instances of gynandromorphism among the swallowtails are rare. Schmid (19 73)-Can.
Entomol. 105: 1549-1551) describes natural gynandromorphs of Ornithoptera victoriae
Gray and O. priamus L. Skinner (1919, Entomol. News 30: 247) and Cockayne
(1935, Trans. Roy. Entomol. Soc. Lond. 83: 509-522) refer to Papilio glaucus L.
gynandromorphs. Hybrid crosses between P. polyxenes and other swallowtails in the
machaon group have in some cases yielded gynandromorphic individuals (Clark and
Sheppard, 1953, Suppl. Entomol. Rec. 65: 1-12; Ae 1964, Bull. Jap. Entomol. Acad.
l: 1-10). Edwards (1868-1872, The Butterflies of North America; Philadelphia:

VoLUME 32, NUMBER 2 139

Am. Entomol. Soc.) presents a figure of an apparently gynandromorphic P. polyxenes,
but offers no data on the specimen.

It is not known what caused the butterfly described here to be a gynandromorph.
Gardiner (1972, J. Res. Lep. 11: 129-140) notes that the incidence of gynandromor-
phism in Pieris brassicae L. cultures is associated with outbreaks of virus and suggests
that viral disease may cause such genetic abnormalities. Viral disease is commonly
present at low levels in our cultures of P. polyxenes and may account for the appear-
ance of this unusual individual.

The specimen is located in Lot 1062 of the Entomological Collections at Cornell
University. I wish to acknowledge the support of N.S.F. Grant DEB 76-20114 (to
Paul P. Feeny ) which covered costs of publication.

Witt1uaM S. Biau, Department of Entomology, Cornell University, Ithaca, New
York 14853.

Journal of the Lepidopterists’ Society
32(2), 1978, 139-140

ELECTROSTRYMON ANGELIA ANGELIA (LYCAENIDAE):
THE OLDEST FLORIDA RECORD?

The butterflies, Sphingidae and Castniidae from the Strecker collection were trans-
ferred from the Field Museum of Natural History to this institution in 1976 on semi-
permanent loan in order that they might be more readily utilized by students of these
groups. Since that time workers here and elsewhere have made greater use of the
Strecker and Reakirt material contained in Strecker’s collection.

That collection is, however, an aggravating one by modern standards: there are
few labels on individual specimens; rather, the series are labelled with data that in
theory apply to all members under the label. This is certainly not the case in all
instances. Fortunately, Strecker prepared catalogs to the Papilionidae, Pieridae and
Lycaenidae before his death, and in these families the data for individual specimens
are recorded.

A second problem involves Strecker’s apparently too-eager interpretation of what
specimen was what, and from where. Some of the putative Reakirt types in the
collection may not be those, and we suspect that Strecker was easy prey for dealers
who peddled material mislabelled by locality. The situation with the Strecker col-
lection is by no means as bad as that with some other older collections, notably the
Ehrmann collection presently housed at Carnegie Museum of Natural History.

Nevertheless, the Strecker collection contains some magnificent material—material
that is not duplicated in other North American collections. Further, the Strecker
collection, with its associated letters, is an historical document. The Lycaenidae,
because that family was one for which the catalog was completed, are especially inter-
esting. In working through the hairstreaks in the Strecker collection, specimens of
Electrostrymon angelia (Hewitson) were found under the label “Thecla hugon
Godart’, a synonym of Electrostrymon endymion (Fabricius ). Two of these specimens
were placed in the collection after Strecker had compiled the catalog and are indi-
vidually labelled “Haiti” and “Port au Prince, Haiti’; both of these butterflies are
specimens of the Hispaniolan subspecies boyeri (W. P. Comstock and Huntington).
The other specimen is labelled characteristically with an “a”, referring to an entry in
the catalog. This specimen is here figured (Fig. 1) and is referable to the Cuban
E. a. angelia. The catalog states that the specimen was from “Florida” and that

140 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fig. 1. Electrostrymon angelia angelia (Hewitson), ¢ upper (left) and under
(right) surfaces; FLORIDA, Chas. Dury (Strecker collection). Allyn Museum photos
071477-15/16.

Strecker received it from Charles Dury. No date is given, but it is likely that the
specimen was taken before 1880.

Kimball (1965, Lepidoptera of Florida) does not list Dury among the pioneer
Florida collectors, and it is possible that while Strecker received the angelia from
Dury, Dury himself may not have collected it. Were it not for the fact that Anderson
(1974, J. Lepid. Soc., 28: 354-358) had recorded this species from the Florida Keys,
and others have reported it from as far north as the Fort Lauderdale area on the east
coast of the state, it would be tempting to dismiss the Strecker specimen as a hoax or a
mislabelled specimen. {! suspect, though, that the Dury/Strecker specimen is an
authentic one, and quite possibly angelia long has been a member of the Florida
fauna, though perhaps not so commonly as in the past few years.

It is further likely that specimens of angelia may have been responsible for the long-
standing records of E. endymion from Florida. The specimen that Holland (1931,
The Butterfly Book: pl. 64, Fig. 32) figured as “endymion” was actually a specimen
of E. angelia boyeri (Klots, 1951, Field Guide to the Butterflies ...: 281), and the
latter author expressed doubt about the occurrence of endymion in Florida. Riley
(1975, Field Guide . . . Butterflies of the West Indies) does not mention endymion

from the West Indies, thus strongly suggesting that the species never has occurred
in Florida.

Lee D. Mitter, Allyn Museum of Entomology, 3701 Bay Shore Road, Sarasota,
Florida 33580.

Journal of the Lepidopterists’ Society
32(2), 1978, 140-141

OBSERVATIONS ON ERORA LAETA (LYCAENIDAE)
IN NEW HAMPSHIRE

Erora laeta Edwards, often considered the rarest of eastern butterflies, is eagerly
sought by many lepidopterists; too often with negative results. I made my first trip
to New Hampshire to capture this species on 21 May, 1977 with fellow collector
Reginald Webster. We visited a few areas in Carroll County, near Bartlett, in
northern New Hampshire, where R. W. had taken one specimen the previous year.

VoLUME 32, NUMBER 2 141

The location seemed typical for /aeta—an abandoned dirt road going through a
beech woods.

We hiked up the road until we reached a shallow gulley crossing the road. On the
other side of the gulley, on damp dirt, was a female E. laeta, which was quickly
netted. We then saw another female which was also easily caught. At this point, I
was amazed, for we had hoped to find one or two E. laeta in a day of intense collecting,
and had caught two in 10 minutes. Continuing along the road, we came to a little
trickle of a stream where we caught two more laeta, both females. Upon returning
to the first gulley, we caught three more females.

Astounded by our luck we decided to try a few more dirt roads in the area. On
every one, we found laeta. It was incredible; was this really that rarest of butterflies
that we were seeing everywhere we looked? In about three hours of collecting we
must have seen over 80 E. laeta, and had collected one or two from every road we
tried, ending up with about 12 specimens apiece. When these numbers are com-
pared to those reported in previous literature (e.g., Mousley, 1923, The Can.
Entomol. 55: 26—29; Field, 1941, Ann. Entomol. Soc. of America 34: 303-316; Smith,
1960, J. Lepid. Soc. 14: 239-240; Roever, 1962, J. Lepid. Soc. 16: 1-4), our sightings
seem truly phenomenal. The most specimens previously reported collected at one time
were two males and nine females along the slopes of Mount Killington in New
Hampshire (Field, op. cit.), and many reports are of individual specimens taken by
chance (e.g., Sullivan, 1971, J. Lepid. Soc. 25: 295-296).

Of all the individuals that we saw, only two were males, only one of which was
captured. Among the females, some were quite worn, while others looked freshly
emerged. There seem two likely reasons, not necessarily mutually exclusive, for the
dearth of males: 1) the males had emerged earlier in the season and so most had
already died, and 2) the males remained up in the trees, and only females came
down to drink at the mud. However the male that was caught was freshly emerged,
and in most species of butterflies, it is the males that are found “puddling’” (Downes,
1973, J. Lepid. Soc. 27: 89-99). The single male was caught in a field next to the
woods.

The females were very easy to catch, some not even flying up when the net was
clapped over them. This behavior has been noted by previous authors (e.g., Hessel,
1952, J. Lepid. Soc. 6:34), but strongly impressed me when I nearly stepped on one
female as I was walking down the road; it flew up from right under my foot. The
males, on the other hand, were more restless and difficult to catch, their flight being
fast and uneven, with only occasional landings on vegetation.

Two possible explanations for the extraordinary abundance of E. laeta that we
observed are as follows: It is possible that there was a real population explosion of
E. laeta in New Hampshire in 1977 (W. Kiel, who has collected for years in New
Hampshire, caught his first E. laeta this spring). It would be interesting to know if
other lepidopterists found a similar increase in this species in other areas. An alter-
native explanation is that /aeta is really not that rare, but that its behavior on this day
was unusual. Perhaps the butterflies normally spend most of their time in the forest
canopy, and thus are not accessible to collectors. This day being very sunny, hot
(33°C), and humid, perhaps drove them down to the ground to drink. It may be that
early emergence and a short flight period make E. laeta seem very rare, but that
finding them is really a matter of being in the right place at the right time. At any
rate, we were!

DEANE Bowers, Dept. of Zoology, University of Massachusetts, Amherst, Massa-
chusetts 01003.

Journal of the Lepidopterists’ Society
32(2), 1978, 142-144

OBITUARY

&

WILBUR S. McALPINE (1888-1977 )

Mr. Wilbur S. McAlpine, charter member of The Lepidopterists’ Society, passed
away on 30 July 1977, at the Grovecrest Convalescent Home in Pontiac, Michigan,
at the age of 88. ‘Mac,’ as he was known by many of his friends, deeply enjoyed
nature and the out-of-doors, and was a devoted amateur lepidopterist, spending most
of his spare time collecting and studying Michigan moths and butterflies. He was
particularly interested in the life history of many local Oakland County species and
specialized in the genus Calephelis, the metalmark butterflies.

Wilbur was born in Detroit, Michigan, on 30 December 1888, and graduated
from Detroit Central High School in January, 1908. He held positions as a draftsman
with the U.S. Lake Survey, Michigan Central Railroad, and Detroit Edison, and was
also employed as an Assistant Surveyor of Coal Claims in Homer, Alaska during
1906, 1907, 1911 and 1912. He served with the military in the 472nd Engineers
during the First World War in 1918. Wilbur became the principal and owner of
a mapping, surveying and engineering business under the names of McAlpine
Engineers, Inc. and W. S. McAlpine Map Co. from December, 1915, until his retire-
ment in 1965, when he sold his business to the employees. They are still conducting
the business under the same names.

While operating his business, the firm produced complete maps of all Michigan
counties, especially detailed maps of Oakland County. McAlpine published an Atlas
of Oakland County, and engineered and recorded over two hundred subdivision plats
and made numerous farm, residential lot and topographic surveys, which are still in
use today. Undoubtedly, many of the Oakland County lepidoptera collected by
McAlpine were discovered during his surveying activities. He witnessed the dis-
appearance of many of his favorite collecting sites due to the suburban encroachment
moving Outward from Detroit.

McAlpine will always be remembered by mid-western lepidopterists as the one who
described the Swamp Metalmark, C. muticum, and subsequently worked out its life
history. In 1971, he culminated his intense interest in the Calephelis with his publica-
tion On the revision of the genus, describing 25 new species and 7 new subspecies

VoLUME 32, NUMBER 2 143

mostly from Mexico and Central America. McAlpine also worked out the life history
of several other Michigan species, including Hyalophora columbia (Smith), Callophrys
and Hesperiidae species, which unfortunately were never published. He was one of
the first to locate several H. columbia tamarack bogs in southeastern Michigan, and
eventually secured numerous cocoons and reared many in his backyard cages.
McAlpine was a very determined Michigan butterfly collector who would travel
anywhere, anytime to add a new species to his collection. Unfortunately, the many
years devoted to his Calephelis project left him with little time for his personal
collection and other lepidopteral plans and pursuits. One of his plans that failed to
materialize due to his death was the publication of a guidebook of Michigan butter-
flies, complete with colored plates of all known species found in the state. This project
was a lifelong ambition to stimulate appreciation for and further the knowledge of
Michigan butterflies, especially among young people. Two of his friends of long
standing, Dr. George W. Rawson and John H. Newman, and this writer to serve
as editor, were to collaborate with ‘Mac’. Although this project is still continuing, the
extremely high cost of colored plates has made it necessary to re-adjust the original
goal.

In 1972, McAlpine donated the bulk of his collection, over 12,000 specimens, in-
cluding many Calephelis type specimens, to the Smithsonian Institution. Later, he
also donated approximately 800 moths and butterflies and 700 miscellaneous insects
to the collection at Michigan State University at East Lansing, and a lesser amount
to The University of Michigan at Ann Arbor. His collection, rich in Michigan material,
included a long series of H. columbia and its cecropia hybrid, Colias interior (Scudder )
and Oeneis chryxus strigulosa (McDunnough ), and miscellaneous Alaskan lepidoptera
and other insects.

McAlpine was an Honorary Member of The Michigan Entomological Society and
held memberships in the former Detroit Entomological Society, The Entomological
Society of Canada and The Lepidoptera Research Foundation, Inc. He was affiliated
with the Cranbrook Institute of Science in Bloomfield Hills, and was made a Life
Member of The Michigan Society of Registered Land Surveyors on 11 February 1970.
He travelled widely in the United States and Mexico and examined museum collections
and collected Calephelis material; and also visited the British and Paris museums in
connection with his metalmark studies.

In addition to his interest in lepidoptera, ‘Mac’ was a devoted and active churchman.
He particularly enjoyed evangelistic singing as a soloist, and sang in the choir at the
First Baptist Church of Birmingham. He and his late wife, the former Minnie
Burnett, are survived by one son, Wilbur Burnett, plus 16 nieces and nephews. Mrs.
McAlpine died in January, 1975.

I wish to express my appreciation to Wilbur's brother-in-law, Mr. Percy C. Burnett
of Pontiac, and his minister, Dr. Glenn H. Asquith, Jr. for information in preparing this
manuscript.

BIBLIOGRAPHY OF WILBUR S. MCALPINE

McAtpine, W. S. 1918. A Collection of Lepidoptera from Whitefish Point, Mich-
igan. Occ. Papers Mus. Zool. Univ. Mich. 54: 1-26, 1 folding map.

1936. Habitat of Cissia mitchellii in Cass County, Michigan. Bull. Brook-

lyn Entomol. Soc. 31: 110, 221.

1937. A Case of Mistaken Identity and Discovery of a New Metalmark

(Calephelis) from Michigan. Bull. Brooklyn Entomol. Soc. 32: 43—49, 1 pl.

1938. Life History of Calephelis muticum (McAlpine), Lepidoptera. Bull.

Brooklyn Entomol. Soc. 33: 111-121, 1 pl.

1939. A New Metalmark (Calephelis) from Texas (Lepidoptera, Riodin-

idae). Bull. Brooklyn Entomol. Soc. 34: 75-80, 1 pl.

144 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

_§. P. Huspety AnpD T. E. Puisxe. 1960. The Distribution, Habits, and Life
History of Euptychia mitchellii (Satyridae). J. Lepid. Soc. 14(4): 209-236, 3 pl.
1971. A Revision of the Butterfly Genus Calephelis (Riodinidae). J. Res.
Lepid. 10(1): 1-125, 8 pl.

1972(1973). Observations on Life History of Oarisma powesheik (Parker )
1870. J. Res. Lepid. 11(2): 83-93, 2 pl.

M. C. Nietsen, Adjunct Curator, Department of Entomology, Michigan State
University, East Lansing, Michigan 48824.

Journal of the Lepidopterists’ Society
32(2), 1978, 144

BOOK REVIEW

FREDERICK WiLLIAM FRoHAWK, by Valezina Bolingbroke, 1977. E. W. Classey Ltd.
Park Rd. Faringdon, Oxon., England. 16 p., 3 figures (photographs), cover drawing
in color by F.W.F.

“<

... 1am going to attempt to give just a brief ‘impression’ of him,” writes the
author on page 1 of this little memoir. The author is not an artist nor a writer as
was Mr. Frohawk, nor a fellow naturalist, but his own daughter. It is obvious from
the start that she felt for him the love and admiration which any father hopes to inspire
in his daughter. The narrative is a series of glimpses into the past, rather like im-
pressionist paintings seen from a distance, as Ms. Bolingbroke rambles through the
English countryside with her readers. One sees an ancient house with moat and
drawbridge, a little stone village, an old fashioned garden, a field of cowslips, travellers
in a ‘pony trap’—all interspersed with charming and humorous incidents, flashes into
the character of her father, his colleagues, his accomplishments and his deep love for -
all of nature, from a very small butterfly to a very young human being. Frohawk
was both author and illustrator of distinguished books on Ornithology and Lepidopter-
ology, but his vast knowledge encompassed many other aspects of nature including
wild flowers, reptiles and weather patterns, to name but a few. One is left wanting
to know much more than this ‘brief impression’ gives us of F.W.F., as Frohawk
was affectionately nicknamed by his friends.

Jo Brewer, Editor, The News of the Lepidopterists’ Society, 257 Common St.,
Dedham, MA 02026.

EDITORIAL STAFF OF THE JOURNAL
AusTIN P. Puatr, Editor

Department of Biological Sciences
University of Maryland Baltimore County, 5401 Wilkens Avenue
Catonsville, Maryland 21228 U.S.A.
Dovuctas C,. Fercuson, Associate Editor THEODORE D. SARGENT, Associate Editor

NOTICE TO CONTRIBUTORS

Contributions to the Journal may deal with any aspect of the collection and study
of Lepidoptera. Contributors should prepare manuscripts according to the following
instructions.

Abstract: A brief abstract should precede the text of all articles.

Text: Manuscripts should be submitted in duplicate, and must be typewritten,
entirely double-spaced, employing wide margins, on one side only of white, 8% x 11
inch paper. Titles should be explicit and descriptive of the article’s content, including
the family name of the subject, but must be kept as short as possible. The first men-
tion of a plant or animal in the text should include the full scientific name, with
authors of zoological names. Insect measurements should be given in metric units;
times should be given in terms of the 24-hour clock (e.g. 0930, not 9:30 AM).
Underline only where italics are intended. References to footnotes should be num-
bered consecutively, and the footnotes typed on a separate sheet.

Literature Cited: References in the text of articles should be given as, Sheppard
(1959) or (Sheppard, 1959, 196la, 1961b) and all must be listed alphabetically
under the heading LirerAtTure Crrep, in the following format:

SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd. ed. Hutchinson,
London. 209 p.

196la. Some contributions to population genetics resulting from the
study of the Lepidoptera. Adv. Genet. 10: 165-216.

In the case of general notes, references should be given in the text as, Sheppard
(1961, Adv. Genet. 10: 165-216) or (Sheppard 1961, Sym. Roy. Entomol. Soc.
London 1: 23-30).

Illustrations: All photographs and drawings should be mounted on stiff, white
backing, arranged in the desired format, allowing (with particular regard to lettering )
for reduction to their final width (usually 4% inches). Illustrations larger than 8%
x 11 inches are not acceptable and should be reduced photographically to that size
or smaller. The author’s name, figure numbers as cited in the text, and an indication
of the article’s title should be printed on the back of each mounted plate. Figures,
both line drawings and halftones (photographs), should be numbered consecutively
in Arabic numerals. The term “plate” should not be employed. Figure legends must
be typewritten, double-spaced, on a separate sheet (not attached to the illustrations),
headed EXPLANATION OF FicuREs, with a separate paragraph devoted to each page of
illustrations.

Tables: Tables should be numbered consecutively in Arabic numerals. Headings
for tables should not be capitalized. Tabular material should be kept to a minimum
and must be typed on separate sheets, and placed following the main text, with the
approximate desired position indicated in the text. Vertical rules should be avoided.

Proofs: The edited manuscript and galley proofs will be mailed to the author for
correction of printer’s errors. Excessive author’s changes at this time will be charged
to authors at the rate of 75¢ per line. A purchase order for reprints will accompany
the proofs.

Correspondence: Address all matters relating to the Journal to the editor. Short
manuscripts such as new state records, current events, and notices should be sent to
the editor of the News: Jo Brewer, 257 Common Street, Dedham, Massachusetts
02026 U.S.A.

ALLEN PRESS, INC. eRINTED LAWRENCE, KANSAS
US.

CONTENTS

STUDIES ON THE INTERACTIONS OF MORPHO PELEIDES (MORPHIDAE)
with LecuMINosAE. Allen M. Young ae |

Notes AND DESCRIPTIONS OF EUPTYCHINI (LEPIDOPTERA: SATYRIDAE)
FROM THE Mexican Recion. Lee D. Miller
LARVAL FoopPpLANT, Lire History NOTEs AND TEMPORAL DISTRIBU-
TION FOR SPLENDEUPTYCHIA KENDALLI (SATYRIDAE) FROM
Mexico. Roy O. Kendall:
Notes ON THE LirE CycLeE AND NATURAL History oF VANESSA
ANNABELLA (NYMPHALIDAE). Thomas E. Dimock

A New Spectres oF HEMILEUCA FROM THE SOUTHWESTERN UNITED

STaTEs (SATURNUDAE). Paul M. Tuskes __... ee |

THe STATUS OF OLLIA PARVELLA DyArR: REDESCRIPTION OF IT IN A
New Genus (Pyratmae). André Blanchard _

Bionomic NOTES ON THE BLOOD-SPOT SKIPPER | HESPERIIDAE: PHOCIDES
LILEA SANGUINEA (SCUDDER)]. Raymond W. Neck __

CiimaTic ReEcIMES RESULTING IN UNUSUAL OCCURRENCES OF
RHOPALOCERA IN CENTRAL TExAs IN 1968. Raymond W. Neck

A New NAME FoR PAPILIO CERES CRAMER, [1776], Nec Fasrictus,
[1775] (NymMpHALmpAE, DANAINAE). Gerardo Lamas _....

Two New PINE-FEEDING SPECIES OF COLEOTECHNITES (GELECHUDAE).
Ronald W. Hodges and Robert E. Stevens _......

Lire History AND HaBits OF COLEOTECHNITES EDULICOLA (GELECHI-
IDAE) A Pinyon NEEDLE MINER IN THE SouTHWEsST. Robert E.
Stevens, J. Wayne Brewer, and Daniel T. Jennings _...

Meyrick’s Recorp oF “MECYNA FURNACALIS, GN.” FROM FIJI, WITH
A New GENERIC ASSIGNMENT FOR PYRAUSTA HOMALOXANTHA
Meyrick (PyYRALIDAE: PyRAUSTINAE). Eugene Munroe and
Akira Mutuura

“ORANGE” BANDs, A SIMPLE RECESSIVE IN ANARTIA FATIMA (NYM-
PHALIDAE). Annette Aiello and Robert E. Silberglied
GENERAL NOTES

Pieris Hadi oleracea (Pieridae) caught by insectivorous plant. Frances S.
Chew i en
Screech owl preys on Peridoma plecta (Noctuidae). Dwight G. Smith _.
A gynandromorph of Papilio polyxenes (Papilionidae). William S. Blau ..
Electrostrymon angelia angelia (Lycaenidae): The oldest Florida record?
Lee D. Miller — a

Observations on Erora laeta (Lycaenidae) in New Hampshire. Deane
Bowers

OBITUARY

Se a HS Hf se fe Si eo

Book REVIEW

65

75

86
88

97

103

107

111

116

118

123

130

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> te fe — Pies = he
ee Sn gee

ort.

Volume 32 1978 Number 3

JOURNAL

of the

LEPIDOPTERISTS’ SOCIETY

Published quarterly by THE LEPIDOPTERISTS’ SOCIETY

Publié par LA SOCIETE DES LEPIDOPTERISTES
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN
Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS

14 November 1978

THE LEPIDOPTERISTS’ SOCIETY |

EXECUTIVE COUNCIL

I. F. B. Common, President T. Surrézu, Vice President
C. V. Covet, Jr., Ist Vice President Jut1an P. Donanue, Secretary
L. A. GozmAny, Vice President RONALD LEUSCHNER, Treasurer

Members at large:

R. A. ARNOLD J. F. EMMEL C. D. FERRIs
E. D. CAsHATT R. R. GATRELLE J. Y. MILLER
R. E. STANFORD AE) Paar M. C. NIELSEN

The object of the Lepidopterists’ Society, which was formed in May, 1947 and
formally constituted in December, 1950, is “to promote the science of lepidopterology
in all its branches, ... . to issue a periodical and cther publications on Lepidoptera,
to facilitate the exchange of specimens and ideas by both the professional worker and
the amateur in the field; to secure cooperation in all measures” directed towards
these aims.

Membership in the Society is open to all persons interested in the study of
Lepidoptera. All members receive the Journal and the News of the Lepidopterists’
Society. Institutions may subscribe to the Journal but may not become members.
Prospective members should send to the Treasurer full dues for the current year,
together with their full name, address, and special lepidopterological interests. In
alternate years a list of members of the Society is issued, with addresses and special
interests. There are four numbers in each volume of the Journal, scheduled for
February, May, August and November, and six numbers of the News each year.

Active members—annual dues $13.00
Student members—annual dues $10.00
Sustaining members—annual dues $20.00
Life members—single sum $250.00
Institutional subscriptions—annual $18.00

Send remittances, payable to The Lepidopterists’ Society, and address changes to:
Ronald Leuschner, 1900 John St., Manhattan Beach, California 90266 U.S.A.

Back issues of the Journal of the Lepidopterists’ Society, the Commemorative
Volume, and recent issues of the NEWS are available from the Assistant Treasurer.

The Journal is $13 per volume, the Commemorative Volume, $6; and the NEWS,
$.25 per issue.

Order: Mail to Charles V. Covell, Jr., Memoirs Editor, Department of Biology, Uni-
versity of Louisville, Louisville, KY 40208, U.S.A.

The Lepidopterists’ Society is a non-profit, scientific organization. The known
office of publication is 1041 New Hampshire St., Lawrence, Kansas 66044. Second
class postage paid at Lawrence, Kansas, U.S.A. 66044.

Cover illustration: Dasychira dorsipennata larva, dorsal and lateral views. From
Fascicle 22.2, “Lymantriidae,” by Douglas C. Ferguson, in Moths of America North
of Mexico. The drawing was done by E. R. Hodges, Scientific Illustrator, Department
of Entomology, Smithsonian Institution. (Reproduced by permission of the author.)

J OURNAL OF

Tue Leprporprerists’ SOCIETY

Volume 32 1978 Number 3

Journal of the Lepidopterists’ Society
32(3), 1978, 145-159

THE INFLUENCE OF ENVIRONMENTAL FACTORS ON
ROOSTING IN THE BLACK SWALLOWTAIL, PAPILIO
POLYXENES ASTERIUS STOLL (PAPILIONIDAE)

JoHN Epwarp Raw ins! AND RoBERT C. LEDERHOUSE?

ABSTRACT. Black swallowtails, Papilio polyxenes asterius Stoll, roost singly in
west sloping old-fields in central New York. A frenetic search flight, triggered by
decreasing radiation regardless of temperature, precedes roosting. Search flights allow
swallowtails to check the suitability of various roosts. Roosts selected favor dorsal
basking for as long as possible under decreasing solar radiation. Dorsal basking on the
roost ceases, and a roosting posture is assumed when a body temperature high enough
for flight can no longer be maintained. Selection for efficient roosting is strong since
most adult deaths apparently occur when individuals are roosting. Selection for
mimetic or cryptic wing patterns is expected to influence the exposed undersurface of
the hindwing more strongly than other wing surfaces hidden during roosting.

Butterflies are active during daylight hours when ambient conditions
of radiation and temperature are high enough to allow behavioral
maintenance of body temperatures suitable for flight (Vielmetter, 1958;
Watt, 1968; Heinrich, 1972). As radiation and ambient temperature fall
during the evening, butterflies seek night roosts where they remain until
the next morning. Some species roost in aggregations (Crane, 1957;
Urquhart, 1960; Benson and Emmel, 1973; Larsen, 1973; Muyshondt and
Muyshondt, 1974; Turner, 1975; Young and Thomason, 1975), but most
roost singly.

Little is known of roosting behavior in most butterflies in spite of the
widely read query concerning roosts in the introduction to Klots’ (1951:7)
popular field guide. The popular literature is full of comments about
butterflies disappearing into trees and fields during evening hours but no
detailed study has ever been reported for a species which roosts singly.

1 Department of Entomology, Cornell University, Ithaca, New York 14853.
Section of Neurobiology and Behavior, Cornell University, Ithaca, New York 14853. Current
address: Department of Zoology and Physiology, Rutgers University, Newark, New Jersey 07102.

146 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Many butterflies bask in the sun as long as possible in the evening,
otten on the actual perch that will be used as the night roost. Such
behavior has been termed “vesper warming” by Clench (1966). Basking
on the roost has been recorded in the hesperiids Thymelicus lineola
Ochsenheimer and Ancyloxypha numitor Fabricius, the lycaenids Everes
comyntas Godart and Lycaena phlaeas L., and the nymphaline Phyciodes
tharos Drury (Clench, 1966). Powell (1968) records similar behavior for
Incisalia iroides (Boisduval) (Lycaenidae).

Several authors have suggested that some species choose roosts which
are shaded in the setting sun and therefore illuminated by the rising
sun earlier the following morning (MacNeill, 1964; Clench, 1966). Such
behavior would favor earlier warming by basking the following morning
and could conceivably lengthen the period of diurnal activity. Roosting
in shadows cast by the setting sun has been observed in Hesperia spp.
(MacNeill, 1964) and in the satyrines Euptychia rubricata Edwards and
Cercyonis meadii Edwards (Clench, 1966). MacNeill described roosting
Hesperia as flying directly away from the sun and roosting on the eastern
side of bushes. Clench noted the movements of a Speyeria aphrodite F.
(Nymphalidae). By lining a tree up with the setting sun and then
flying directly toward the sun, the butterfly arrived at the shaded side of
the intervening tree. After repeating this behavior three or four times,
it flew off and roosted in a field. It remains unclear if such shade seeking
during roosting is widespread among butterflies, let alone the mechansm
by which this is accomplished.

The environmental cues which release roosting behavior have been
inadequately studied. When ambient radiation and temperature are low
enough to prevent maintenance of a body temperature suitable for flight
in the nymphalid Argynnis paphia (L.), roosting results (Vielmetter,
1958). This does not explain the onset of roosting on warm evenings when
ambient temperatures allow sustained and active flight in the absence of
direct insolation. Larsen (1973) has suggested that the timing of roosting
events may depend more on decreasing radiation from the setting sun
than on decreasing ambient temperature. 4

From the preceding it seems that selection of the roost site in a given
species may involve one of three different roosting patterns: 1) Roosting
in shadows cast by the setting sun, therefore hastening exposure to the
rising sun the following morning, 2) roosting and basking in areas having
maximal exposure to the setting sun, thereby prolonging the period
when body temperatures suitable for flight may be maintained, or 3)
roosting in areas not related to the position of the sun.

The following study examines in detail the roosting behavior of the

VOLUME 32, NUMBER 3 147

black swallowtail, Papilio polyxenes asterius Stoll, and determines what
environmental cues direct such behavior. It constitutes the first study
of roosting in papilionids [see observations of Gibson and Panchen (1975)
on the African Papilio demodocus Esper] and the first detailed study
of non-gregarious roosting for any butterfly.

MATERIALS AND METHODS

The black or parsnip swallowtail, Papilio p. asterius is frequently seen
in open clearings and fields throughout eastern North America. The
territorial males may be observed for lengthy periods in relatively small
areas (Lederhouse, 1978). The darker females are more transient, usually
observed flying quickly across these open areas.

Roosting was studied in an old field near Brooktondale, southeast of
Ithaca, Tompkins County, New York during the summer of 1975 (Fig. 3).
This field was bounded on the north and east by corn fields and on the
south and west by dense woody growth. The field was divided into a
grid consisting of 276 squares each 10 meters on a side. The relative
elevation of each corner of every grid square was established by sur-
veying. Slopes and aspects (compass headings) for every grid square
were determined by treating each square as a planar surface with corner
elevations most nearly matching those measured. Plant densities for
each square were measured by direct count for large, conspicuous species
or by counting within a randomly placed 0.5 m? circle for smaller and
more numerous species. Only the most abundant plant species are treated
in this study. Plant nomenclature follows Fernald (1950).

During each study period, gross solar radiation and ambient tem-
perature were recorded using a centrally-located radiometer and hygro-
thermograph. Sunset times were established by reference to published
readings of the U.S. Naval Observatory for Binghamton, New York
(U.S. Printing Office, 1959).

Observations of individual behavior were made by persistently fol-
lowing one swallowtail during the evening hours until it selected a roost.
This was difficult due to the rapid and erratic flight of individuals seeking
roosts; more often than not the butterfly was lost from sight. Additional
data were obtained by systematically walking over the entire grid during
dusk noting the location and microhabitat of individuals already on
roosts. Observations of behavior during the middle of the day were
taken from concurrent research by one of us (J. E. R.) and used for
comparative purposes.

Clock times (EDT) were recorded at the onset of flights preceding
roosting. Times were also recorded when butterflies were seen to assume

148 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TABLE 1. Behavioral responses and ambient environmental factors associated with
roosting in Papilio polyxenes compared with values for those variables associated with
other diurnal activity.

Roosting Non-roosting
5d est fe! D) (CNY) xt S.D. (N)
Flight duration** (seconds ) 96 + 111 (19) 33 = 52 (651)
Perch duration (seconds ) (Overnight ) 132 + 208 (807)
Perch height* (cm) 58} ae Iles} (8S) 64 + 27 (794)
Air temperature 150 cm above the
ground when roost posture assumed**
( Centigrade ) PPS AS) as B5) (1510) DAES) a= PLT tl0 Ne)
Relative humidity when roost
posture assumed** (7%) 76 + 16 (37) 58 + 11 (640)
Solar radiation when roost posture
assumed** (cal/cm’/min) 16 = .20 (15) Oo = mlian(254))

Asterisks indicate significance level of two-tailed Student’s t-test for differences between roosting
and non-roosting mean values: * (.05 >P > .01); ** (P < .01).

a roosting posture with folded wings or when they were discovered already
in this posture. Identification of individuals at a distance was made pos-
sible by marking the marginal and submarginal row of wing spots with
felt-tip pens (Lederhouse, 1978).

RESULTS

Roosting Search Flight

Roosting was always preceded by an extremely rapid and erratic
flight of longer average duration than flights at other times of the day
(Table 1). The shift from non-roosting activity into this search flight
was not gradual but occurred abruptly, the individuals accelerating and
often disappearing from sight in seconds. In the 8 cases where a single
butterfly was observed throughout a search flight, movement was seen
to be circular, the buttertly flying in circles often more than 5 m above
the ground. During these flights, which avoided shaded areas, individuals
often returned to certain spots in the field, flying low and dipping down
into the vegetation.

Search flights were repeatedly interrupted by short periods of perching,
lasting one or two seconds. These were on potential roosts which either
were not stable in the wind, were deflected greatly by the butterfly’s
weight, did not provide a good gripping surface (smooth grass culms ),
were very far from the ground, or were in the deep shade. Butterflies
did not select the roost directly but only remained on those perches which
were suitable after abandoning many perches which were not.

VOLUME 32, NUMBER 3 149

Behavior on the Roost

Eventually a suitable roost was located and the butterfly positioned
itself so that the frontal plane of its body was perpendicular to the incident
radiation from the setting sun, spread its wings, raised its abdomen
between the spread anal margins of the hindwings, and then remained
motionless (Fig. 1). This basking continued for several minutes, being
longer on warm evenings than on cool or hazy ones.

Basking ceased if the butterfly came under the advancing shadows of
the surrounding vegetation. At the end of basking the abdomen was
suddenly lowered, the wings dorsally appressed and swept back such
that the hindwings almost totally covered the front wings, and the body
brought parallel with the stem or head of the roost plant (Fig. 2). The
posture was maintained throughout the night.

The following morning, basking occurred as soon as direct sun fell on
the roost. Morning basking appeared identical to that occurring in the
evening. Butterflies were seen to rotate about the roost so as to be
perpendicular to the rising sun. Moming basking was never seen earlier
than two hours after sunrise and usually occurred between 0800 and 0900
(EDT) in July and early August.

Effects of Ambient Temperature and Radiation on Roosting

Mean values of ambient temperature, relative humidity, and solar
radiation all differed between roosting and non-roosting situations (Table
1). It was not surprising that the evening hours associated with roosting
had significantly lower radiation, lower temperature, and higher relative
humidity than earlier in the day.

Papilio polyxenes males are incapable of sustained flight with thoracic
temperatures below 24 C (Rawlins, unpublished). This value matches
that of the average air temperature when the roost posture with folded
wings was observed in the evening (23.9 C). Since radiation was very
low at these times, body temperatures would rapidly approach ambient
levels in quiescent individuals. Several times individuals which had been
roosting less than one minute following vigorous flight were observed to
be incapable of sustained flight if disturbed. For example, one female
disturbed under .09 cal/cm?/min radiation and 23.3 C ambient tem-
perature was unable to remain airborne and crashed into the grass after a
gliding flight of about 5 m. This suggests that the roost posture is assumed
under decreasing radiation when ambient temperatures drop below the
range of body temperatures allowing activity.

The onset of roost search flight during July and August paralleled the
time of meteorological sunset (Fig. 7). This suggests that the onset of

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VoLUME 32, NUMBER 3 tom

TABLE 2. Plant species used as roosts by Papilio polyxenes.

Number on heads Number on stems

Roost species or apices or culms Total
Tragopogon pratensis L.* 4 J. 5
Lonicera morrowi Gray 0 1
Potentilla recta L.* 1! 0 if
Solidago altissima L. 0 2 2
Linaria vulgaris Hill 0 1 1
Total dicotyledons = 5 10
Dactylis glomerata L. 2 0 2
Phleum pratense L. 12 4 16
Other grasses 2 b 9
Total grasses 16 RE DT
Total roosts 21 16 Sil
Percent of total 57% 43% 100%

* All roosts were on dead, leafless scapes of the previous season.

the search flight is cued by decreasing radiation even though flying or
basking on the roost continues until ambient temperatures fall to levels
preventing further activity.

The occurrence of abbreviated search flights during sudden radiation
drops preceding thunderstorms provided additional evidence that de-
creasing radiation triggers such search flights. One male patrolling an
area high in the field abruptly switched into erratic search flight behavior
with the sudden darkening before a thunderstorm even though air tem-
perature 1.5 m above the ground was 26.7 C. After being forced down
minutes later by heavy rain, this male assumed the roosting posture in
an air temperature of ,20.6 C. With the return of direct sun, basking
commenced.

Description of the Roost

Seventy-three percent of the plants selected for roosts were grasses,
the remainder being dicotyledonous species (Table 2). Sixty percent of
the roosts on dicots were on leafless scapes of dead plants from the
previous season. Such plants constituted less than five percent of the
total stems within any grid square in the field.

=

Figs. 1-2. Postures of Papilio polyxenes on the roost. 1. Adult female in dorsal
basking posture on culms of grass. 2. Male in folded-wing posture on short head of
Phleum pratense,

152 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

@-female
€-male

Fig. 3. Topography of study area showing location of roosts. Each small grid
square is 10 m square. Numbers in margins give elevation of isoclines in meters above
lowest point in field (northwest corner).

Roosts on plant inflorescences or apices were more frequent than
roosts on culms or stems. Such heads may provide roosts which are easier
to cling to or are less deflected by the butterfly’s weight.

Butterflies on roosts are significantly closer to the ground than those
perched during daily activity (Table 1). The maximum height of vegeta-
tion within a one meter radius of each roost averaged greater than twice
the roost height (110 + 17 cm, N = 33). Roosts were invariably in open
areas of the vegetation where contact of the roosting butterfly with sur-
rounding plants was impossible in all but the heaviest winds.

Description of the Roost Sites

Roosting occurred primarily on sites which sloped toward the west
and were at the higher elevations in the field (Fig. 3). There were no
differences between the roosting sites of males and those of females.

The distribution of slopes for the ten meter grid squares in which
roosting occurred differed significantly from that of slopes for all grid
squares in the field (Kolmogoroy-Smirnov Test; .05 > P > .01) (Siegel,
1954) (Fig. 4). The average slope of 53 roost sites was 4.2 + 2.5 (S.D.)
degrees above the horizontal. Similarly, the distribution of aspect values
toward which roost sites sloped differed significantly from that for all grid

VoLUME 32, NUMBER 3 153

SOF a ALL GRID SQUARES

Fig. 4
| GRID SQUARES WITH ROOSTS
258
20
% 15

woe S74" \4i-5 J at

ae in degrees above cb hae

O

(n

ALL GRID SQUARES
25 Fig.5 O

|| GRID SQUARES WITH ROOSTS

<240 240249 250259 26026? 270279 280289 290299 300309 310319 7320

Aspect in compass degrees

%

Figs. 4-5. Frequency distributions of slope (4) and aspect (5) values for all
grid squares in study area compared with distributions for only those grid squares
containing roosts.

sites in the field (P < .01) (Fig. 5) even though the field predominantly
sloped toward the west.

The density distribution of yellow goat’s-beard (Tragopogon pratensis
L.) and rough-fruited cinquefoil (Potentilla recta L.) for roosting sites

154

50

40

80

70

60

50

40

30

20

JOURNAL

0 les 4-7 22 Uf

Stems of Tragopogon and Potentilla , m?

0) l= 20: 20-4040
Stems of Solidago and Aster / mi

I ALL GRID SQUARES

| GRID SQUARES WITH ROOSTS

%

%

OF THE LEPIDOPTERISTS SOCIETY

60

0-39 40-79 80-119 7119

Grass Culms / m?*

40
30

20

10

0-19 20-39 40-59 CO79 77
Culms of Phleum and Dactylis | m

50

40

0 1-19 20-39 40-79 >79
Asclepias Plants / 100 m?

Fig. 6. Frequency distributions of various plant densities for all grid squares in
study area compared with distributions for only those grid squares containing roosts.

differed significantly from that for all sites in the field, the density being
higher for roosting sites (Kolmogorov-Smirnov Test; .05 > P > .O1)

(Fig. 6).

The density distribution of goldenrod (Solidago spp.) and New
England aster (Aster novae-angliae L.) also differed significantly (P
< .01). Roosts did not occur where Solidago and Aster were at highest

density; these species usually form dense stands of plants which contact
each other greatly in the wind.

VoLUME 32, NUMBER 3 155

On the other hand, density distributions of all grass species, of just
orchard grass (Dactylis glomerata L.) and timothy (Phleum pratense L.)
taken together, and of common milkweed (Asclepias syriaca L.) did not
differ significantly (P > .05) between roosting sites and all field sites.

Survivorship on the Roost

Roosting individuals were quiescent and could easily be approached
and caught by hand. Such inability to escape suggested that predation
during roosting might be a major, if not the chief, cause of mortality
in black swallowtail adults.

Twenty-six roosts were visited the following morning before basking
occurred. Two male butterflies were missing from their roosts and were
never seen again in the study area. Probability of death per night on the
roost was estimated as being 2/26 = 0.077 for all butterflies regardless of
sex. Thus, an adult could be expected to live through an average of 8.7
nights assuming no other sources of adult mortality were considered
[(1—.077)®:* = 0.5) ]. If only males were considered the probability of
death on the roost per night was estimated by 2/21 = 0.095. The average
life expectancy of males was through 6.9 nights ignoring other sources
of mortality.

DISCUSSION

Black swallowtails favor roosting sites which will allow them to con-
tinue basking for as long as possible under decreasing radiation. Selection
of relatively open roost sites which slope toward the west favors the
maintenance of body temperatures suitable for continued activity for
as long as possible. There is no indication of any behavior which would
favor warming the following morning. The presence of such behavior
in other butterfly species needs more study.

The only unique behavior associated with roosting is the roost search
flight. It is this flight which provides the mechanism by which a suitable
roost is selected. Repetitive brief perches during this flight allow the
butterfly to test and reject perches which are unstable, in the shade, or are
disturbed by other vegetation. Frenetic activity during the search flight
may provide enough body heat metabolically to allow flight to continue
under conditions which would prevent further activity if the butterfly
were motionless for a short period.

Since the search flight occurs in the sun, roosts which are finally
selected tend to be on sites having the longest and most direct exposure to
the setting sun. Roosting individuals on level sites are more susceptible

156 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

1900

aw! |

1830
=
Y 1800
°o
1730 ae
vaeeees two hours before sunset
0
O-onset of a roost search flight
1700

June July July July August August August Sept.
25 ©) 15 JAS) ©) 15 25 5

Fig. 7. Comparison of the time of onset of roost search flights with time of sunset.
The solid line is a least-squares regression line of onset times with respect to date.

to being shaded by plants between them and the sun than are those on
sloping sites facing the setting sun.

The onset of the erratic roost search flight is apparently triggered by
decreasing radiation regardless of temperature. The shift from dorsal
basking on the roost to a roosting posture with folded wings appears to
occur when ambient temperature and radiation are such that body tem-
peratures suitable for flight can no longer be maintained. There appears
to be no endogenous rhythm controlling roosting. Such behavior is just
as Clearly associated with the onset of unfavorable conditions for flignt
at other times of the day.

Basking on the roost is similar to basking under more favorable con-
ditions. There is no difference between morning and evening, making
Clench’s (1966) categories of “vesper” and “matutinal warming” difficult

VOLUME 32, NUMBER 3 157

to define for P. polyxenes. Basking on the roost may lengthen the period
in the evening when escape from predators or selection of a different
roost are still possible should a predator appear or the roost become
unsuitable.

Selection for roosting behavior and roost preferences appears to be
strong in view of the high mortality recorded in this study. Those
individuals which fail to secure a steady, protected roost risk dislodgement
and predation while cold and helpless at ground level.

In a concurrent study, Lederhouse (1978) estimated the probability of
permanent disappearance per day due to dispersal and death for males
after the first night of adult life to be 0.106. This value for males through
the first night of imaginal life was measured as 0.532. The present estimate
of male mortality on the roost (0.0953 is therefore about equal to that
which Lederhouse ascribes to all causes of death plus dispersal because
all marked butterflies in our study were at least two days old. It is con-
ceivable that the large loss through the first night of adult life may be
attributed to failure to find a suitable roost during that first evening or to
considerable dispersal during the first day.

Death on the roost may be the chief source of mortality in this popula-
tion, since not one instance of diurnal predation has been seen in over 700
hours of close observation, notwithstanding reported cases of diurnal
predation by passerine birds (Erickson, 1973). Predation by ants may
explain the preference shown for dead dicot scapes as roosts since ants
may not frequent such plants during foraging.

If death on the roost is the major source of mortality in adult black
swallowtails, then selection for mimetic or cryptic coloration patterns
would probably operate most strongly on that portion of the butterfly
which is exposed during roosting, the undersurface of the hindwings. We
cannot help but note that the undersurface of the hindwings of members
of the supposed Battus philenor mimetic complex are more similar in
coloration than any other part of the wings. This complex includes
B. philenor (L.), Papilio troilus L., P. glaucus L. (dark females), P.
polyxenes F., and Limenitis arthemis astyanax F. (Brower, 1958). We
suggest that selective advantages gained by cryptic or mimetic coloration
patterns in many butterfly species may be greatest during roosting and
that the exposed undersurface of the wing is more affected by such
selection than is the hidden uppersurface.

In conclusion, black swallowtails select roosts which maximize exposure
to the setting sun and provide stable, secure sites preventing dislodge-
ment during the night. A roost search flight triggered by decreasing
radiation is the only behavior unique to roosting and provides a

158 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

mechanism by which suitable roosts may be selected. Selection for
efficient roosting is strong since death on the roost is likely to be the
major source of mortality in adults.

ACKNOWLEDGMENTS

We wish to express our appreciation to Owen Sholes, Dr. Paul Feeny,
and Dr. John G. Franclemont for comments on the manuscript. We thank
Dr. Mark Scriber for suggestions on the implications of roost mortality on
selection for wing coloration.

LITERATURE CITED

Benson, W. W. & T. C. Emme. 1973. Demography of gregariously roosting
populations of the nymphaline butterfly Marpesia berania in Costa Rica. Ecology
54: 326-335.

Brower, J. V. Z. 1958. Experimental studies of mimicry in some North American
butterflies. Part II. Battus philenor and Papilio troilus, P. polyxenes and P.
glaucus. Evolution 12: 123-136.

Ciencu, H. K. 1966. Behavioral thermoregulation in butterflies. Ecology 47:
1021-1034.

Crane, J. 1957. Imaginal behavior in butterflies of the family Heliconiidae:
changing social patterns and irrelevant actions. Zoologica (New York) 42:
135-145.

Erickson, J. M. 1973. Bird predation on Papilio polyxenes F. (Papilionidae). J.
Lepid. Soc. 27: 16

FERNALD, M. L. 1971. Gray’s Manual of Botany. Eighth ed. D. Van Nostrand,
New York. 1632 p.

Gipson, D. O. & A. L. Pancuen. 1975. Roosting behavior of the butterfly Papilio
demodocus Esp. on the Kenya coast. Entomol. Rec. 87: 156-158.

Hernricu, B. 1972. Thoracic temperatures of butterflies in the field near the
equator. Comp. Biochem. Physiol. 42A: 459-468.

Kiors, A. B. 1951. A Field Guide to the Butterflies of North America, East of the
Great Plains. Houghton Mifflin, Boston. 349 p.

Larsen, T. B. 1973. Communal roosting among butterflies in the Lebanon.
Entomol. Scand. 4: 299-301.

LepErRHOUsSE, R. C. 1978. ‘Territorial behavior and reproductive ecology of the
black swallowtail butterfly, Papilio polyxenes asterius Stoll. Cornell University
Ph.D. Thesis. Ithaca, New York.

MacNemz, C. D. 1964. The skippers of the genus Hesperia in western North
America, with special reference to California (Lepidoptera: Hesperiidae). U.
Calif. Publ. Entomol., 35. 221 p. |

Muysnonpt, A. & A. MuysHonptr, Jr. 1974. Gregarious seasonal roosting of
Smyrna karwinskii adults in E] Salvador (Nymphalidae). J. Lepid. Soc. 28:
224-229,

PoweELx, J. A. 1968. A study of area occupation and mating behavior in Incisalia
iroides (Lepidoptera: Lycaenidae). New York Entomol. Soc. 76; 47-57.

SieceEL, S. 1956. Nonparametric Statistics for the Behavioral Sciences. McGraw-
Hill, New York. 312 p.

Turner, J. R. G. 1975. Communal roosting in relation to warning colour in two
heliconiine butterflies (Nymphalidae). J. Lepid. Soc. 29: 221-226.

U.S. Pare. Orr. 1959. Sunrise and Sunset at Binghamton, New York. Nautical
Almanac Office, U.S. Naval Observatory, Wash., D.C.

VOLUME 32, NUMBER 3 159

Urounart, F. A. 1960. The Monarch Butterfly. Univ. of Toronto Press. 361 p.

VIELMETTER, W. 1958. Physiologie des verhaltens zur Sonnenstrahlung bei dem
Tagfalter Argynnis paphia L. I. Untersuchungen im Freiland. J. Insect Physiol.
2: 13-37.

Watt, W. B. 1968. Adaptive significance of pigment polymorphisms in Colias
butterflies. I. Variation in melanin pigment in relation to thermoregulation.
Evolution 22: 437-458.

Younc, A. M. & J. H. THomason. 1975. Notes on communal roosting of Heliconius
charitonius (Nymphalidae) in Costa Rica. J. Lepid. Soc. 29: 243-255.

ANNOUNCEMENT

The Xerces Society offers modest research grants to support scientific work which
has some likelihood of contributing to the preservation of terrestrial arthropod popula-
tions as biological entities. XS is an international, non-profit organization dedicated to
the conservation of terrestrial arthropod populations and their habitats. Awards will
usually be a few hundred dollars but may be larger if funds are available. Knowledge-
able young workers and those without formal professional affiliation are encouraged
to apply. For details write to Dr. FRANcIE CuEw, Xerces Grants Committee Chairman,
Department of Biology, Tufts University, Medford, Massachusetts 02155, U.S.A.

Journal of the Lepidopterists’ Society
32(3), 1978, 160-174

NOTES ON THE LIFE CYCLE AND NATURAL HISTORY
OF BUTTERFLIES OF EL SALVADOR. IIC.
SMYRNA BLOMFILDIA AND S. KARWINSKII

(NYMPHALIDAE: COLOBURINI)

ALBERT MuysSHONDT, JR. AND ALBERTO MuysHONDT
101 Avenida Norte #322, San Salvador, E] Salvador

ABSTRACT. Descriptions and photographs of the life histories of Smyrna
blomfildia and S. karwinskii are presented, and the larval foodplants ( Urticaceae) and
the comparative behavioral characteristics of the two species are recorded and dis-
cussed. The present taxonomic placement of S. blomfildia and S. karwinskii is
questioned, and an alternate interpretation is expressed based on the differing degrees
of morphism between the two species. The adaptiveness of polymorphism is explained
relative to human-disturbed habitats in E] Salvador, noting that the monomorphic
S. karwinskii is the most evolved species, but that S. blomfildia has a flexibility to
Overcome adverse conditions because of certain polymorphic characteristics. The
peculiar phenomenon of a divergent evolutionary trend between the early stages
(larvae and pupae) and an advergent trend between the adults of both species
is noted.

This is the second of three papers on the Coloburini (Gynaeciini) of
El] Salvador. Classically, Colobura dirce L. and Historis odius Fab. have
been included with Smyrna in the Coloburini, based, no doubt, on
similarities in adult characteristics. Whether or not these form a natural
complex of related species is left for others to determine, who can compare
the overall characteristics of Smyrna sp., Colobura dirce, and Historis
odius.

We present here a description of the early stages of Smyrna blomfildia
Fab. and S. karwinskii Hibn., records of the larval foodplants, and an
account of the behavior observed in both the immatures and adults. We
have reared S. blomfildia and S. karwinskii using the same techniques
described for the Catonephelini, Charaxini, and Hamadryadini (Muy-
shondt, 1973a, 1974b; Muyshondt & Muyshondt, 1975a) with consistently
uniform results. Specimens preserved in alcohol have been sent to the
American Museum of Natural History, New York City.

Life Cycle
Smyrna blomfildia Smyrna karwinskii

Egg. Almost spherical with flattened Same as S. blomfildia in all respects.
base, light green with 10 whitish vertical

ribs which fade around the micropyle. Ca.

|! mm diameter. Hatches in 5 days.

VOLUME 32, NUMBER 3

Ist instar larva. Head roundish, naked,
shiny black. Body naked, cylindrical,
brownish-green, with transverse rows of
white, shallow, small warts on each seg-
ment. Legs and tips of prolegs dark
brown. Ca. 2 mm when hatched, growing
to 4.5 mm in 2 days.

2nd instar larva. Head shiny black, with
short, knobby, divergent horns, one on
each epicranium, and 8 small, white con-
ical projections across head capsule, under
epicranial horns. Body brown with trans-
verse rows of tiny, forked spines im-
planted on white chalaza. Grows to 8 mm
in 2 days.

3rd instar larva. Head reddish with
thick, short horns (ca. head length) armed
with secondary spines placed in the fol-
lowing order: a basal row of 3, 1 pointing
inwards, 2 anterad; a second row, with 1
pointing caudad, 1 anterad and 1 laterad.
Horns terminate distally in a club with 5
short spines. Around base of each horn
are 7 spines; around the ocelli are 5
smaller spines. Body predominantly dark
brown with some light spots among the
spines, which are placed in the following
positions: Ist thoracic segment (T-1)
with 1 spine subdorsally, 1 subspiracular
spine and 1 pedal spine. T-2 and T-3
with 1 subdorsal scolus with a rosette of 5
spines near tip and 1 vertically; 1 supra-
spiracular scolus with rosette of 4 spines
and 1 vertical; 1 subspiracular spine and 1
pedal spine. Abdominal segments from
A-l1 to A-7 have, in addition, 1 dorsal
scolus with 2 lateral spines and 1 distal,
and behind the subspiracular spine 1
scolus with rosette of lateral spines and 1
distal spine. A-l1, A-2 and A-7 have 1
small ventral spine in line with prolegs.
A-8 has an additional scolus caudad with
a 6-spined rosette. A-9 with only 1 sub-
dorsal scolus directed posterad with rosette
of 5 lateral spines and one distal. A-10
with anal shield and 2 lateral groups of
small spines directed posterad. Grows to
15 mm in 2 days.

4th instar larva. Same as 3rd instar but
body shows various color morphs: 1
dorsally brown with cream dots, rest of
body cream also, where black spiracula

161

Same as S. blomfildia, but light green
with transverse rows of white warts.

Same as S. blomfildia, but body lighter
color and lacking dorsal spines in all but
8th abdominal segment.

Head light brown with longer and thinner
horns than S. blomfildia (1% head
length). Lateral spines of head much
reduced. Horn terminals more clubbed.
Body color basically brown with double
transverse rows of whitish dots on each
segment and a broken stripe of light color
subspiracularly. Spiracula black. Ventral
surface dirty light gray, prolegs beige.
Body with whitish scoli armed with con-
colored black-tipped spines, placed in the
following order: T-1 with subdorsal
group of 3 small spines, then 1 small
supraspiracular spine, 1 small subspirac-
ular spine and 2 small pedal spines. T-2
and T-3: 1 subdorsal scolus, short, with
rosette of 6 lateral spines and 1 distal
spine; 1 supraspiracular scolus with
rosette of 5 lateral spines and 1 distal:
1 subspiracular spine and 2 pedal spines.
Abdominal segments A-1 and A-7 have a
subdorsal scolus with a rosette of 4 lateral
spines and 1 distal; supraspiracular scolus
with rosette of 4 lateral spines and 1
distal; subspiracular scolus with 4 lateral
spines and 1 distal, then two small pedal
spines. A-8 has in addition a heavier
dorsal scolus armed with 8 spines. A-9
has only 1 supraspiracular scolus directed
posterad, with 6 spines. Anal plate on
A-10 surrounded by 6 small spines.
Grows to 15 mm in 2 days.

Head dirty yellow, with some brown
markings frontally. Slender horns, slightly
bent in some individuals, colored light
gray. All head and horn spines white with

162

stand out and all scoli and spines whitish.
Another morph mostly black dorsally,
with double row of whitish dots along
meson. From supraspiracular to ventral
area cream colored. Dorsal scoli white,
subdorsal and supraspiracular scoli black,
the rest whitish. Other morph mostly
greenish-white with black stripes covering
dorsal and subdorsal area, but much
broken by stripes and dots of greenish-
white. All scoli greenish-white with light
spines. Still another morph similar to the
preceding one but with subdorsal and
supraspiracular scoli black. The rest whit-
ish. Grows to 28 mm in 2 to 3 days.

5th instar larva. Same as 4th instar,
growing to 41 mm in 3 to 4 days.

Prepupa. No noticeable change. Hangs
from anal prolegs, body incurved ven-
trally. Lasts 1 day.

Pupa. From light brown to very dark
brown, abdominal segments darker than
the rest, with rows of lighter, shallow
warts: 1 supraspiracularly, 1 subspirac-
ularly. Spiracula inconspicuously brown.
One black spot at either side between
wingcase and thorax. Abdomen rounded
with no sharp angles; slightly incurved
ventrally, shallow depression at the tho-
racic union dorsally. Thorax slightly
keeled to rounded head. Pointed cre-
master dark brown. 25 mm long, 10 mm
laterally and dorsoventrally at widest
points. Adults emerge between 8-11
days.

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

black tips. Body ground color light gray
with a darker thin mesal stripe, and trans-
verse rows of cream colored spots at seg-
ment unions. All scoli as in 3rd instar,
implanted now on bright yellow chalaza;
scoli and spines white with black tips.
Grows to 29-30 mm in 2 to 3 days.

Drastic change in color. Now mostly
brown with black markings on head, be-
tween dark horns and down to ocelli lat-
erally. The whole body with black ir-
regular marblings. All scoli shorter than
in prior instars; brown colored as well as
spines, legs and prolegs. Grows to 40—42
mm in 3 to 4 days.

Same as S. blomfildia. 1 day.

Lighter brown and thinner than S. blom-
fildia. Abdomen more humped dorsally
and with 3 rows of conical spines: 1
prominent subdorsally, 1 of decreasing
size supraspiracularly, and 1 still smaller
subspiracularly. Thoracic dorsal keel
sharply angled midways. Black spots be-
tween wingcase and thorax, as S. blom-
fildia. Black pointed cremaster set at an
angle in relation to body plan. 28 mm
long, 11 mm dorsoventrally, 10 mm lat-
erally at widest points. Lasts 9-11 days.

Adults. Both species show a marked sexual dimorphism in the coloration, males
being brighter than females. Wing shape is the same in both sexes.

Males. Dorsally forewing golden brown in basal and discal areas with lighter golden-
orange band slanting from midcostal margin to tornus. (S. blomfildia tends to be
lighter than S. karwinskii.) The rest velvety black, except a subapical row of three,
light yellow spots parallel to the light band. Hindwing in S. karwinskii golden-brown
mostly, with a black edge along outer margin, 3 mm wide, near outer angle, very thin
from there down along the very edge of the wing to anal angle. A submarginal row of
faint black dots along the thin portion of the black edge. In S. blomfildia hindwing is a
lighter golden-orange and black edge along outer angle becomes submarginal along

VoLUME 32, NUMBER 3 163

Figs. 1-11. Smyrna blomfildia: 1, egg, recently deposited, 1 mm; 2 (photo upside-
down), lst instar larva, 4.5 mm (note “perch” on central vein); 3, 2nd instar larva,
8 mm (note new “perch” being constructed at edge of leaf); 4, 3rd instar larva,
15 mm; 5, 4th instar larva, 28 mm; 6-10, 5th instar larvae, various morphs, 41 mm;
11, 5th instar larva, close-up of head.

outer margin down to Mz where it becomes thin, ending between Cu: and Cuz where
it is substituted by a marginal thin edge running between the two, small toothed
projections on anal angle, with a whitish dot in the interior one. Inner fold in both
species fulvous gray.

Ventrally forewing presents basally some black drawings, on light yellow basic
color, which is devoid of markings from midcostal margin to tornus, except for a
brownish gray border along inner margin, more so in S. karwinskii. From midcostal
to subapical costal margin down to mid-outer margin to tornus there is a black zone

164 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Figs. 12-17. Smyrna karwinskii: 12, egg, ready to hatch, 1 mm; 13, Ist instar
larva, 3.5 mm; 14, 3rd instar larva, 15 mm; 15, 4th instar larva (note absence of
dorsal spines and slender horns ), 29-30 mm; 16, 5th instar larva (note reduced scoli),
40-42 mm; 17, 5th instar larva, close-up of head.

limited distally by a diffuse replica of the dorsal subapical dots, which ventrally merge
with each other. Apically a gray zone mottled by faint black markings, more con-
trasting in S. blomfildia than in S. karwinskii.

Hindwings show a complicated pattern of sinuous lines, circles and triangles of dark
brown, light brown and whitish color, all darker in S. blomfildia than in S. karwinskii.

VoLUME 32, NUMBER 3 165

Figs. 18-20. Smyrna blomfildia: 18, dorsal view of pupa; 19, side view of pupa;
20, ventral view of pupa.

Figs. 21-23. Smyrna karwinskii: 21, dorsal view of pupa; 22, side view of pupa;
23, ventral view of pupa.

166 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Figs. 24-27. Smyrna blomfildia: 24, dorsal view of male; 25, ventral view of male
(metric scale); 26, dorsal view of female; 27, ventral view of female.

Both species present a submargnal row of 4 “eyes” along outer margin, the two at the
extremes twice as large as the two interior ones. Both species also have a black spot
on the anal angles. S. blomfildia in addition has a second black spot on the first
toothed projection.

Females. Both species dorsally have the same pattern as the males, but the golden-
brown or orange is replaced by dull brown, separated from the black apical area by a
light yellow band. The rest as in males and so is the underside of the wings.

Body is concolorous to the respective wing coloration. Antennae black, ending with
an orange tip, larger in S. blomfildia. Palpi cream colored, proboscis tanned brown.
Wing span 70-80 mm. S. blomfildia usually larger than S. karwinskii, and females
larger than males, more markedly so in S. blomfildia.

Total time from egg to adult, 25-30 days.

Natural History

Females of both species of Smyrna deposit their solitary eggs on the
undersides of leaves of various Urticaceae. In contrast, other species of
Coloburini oviposit on Moraceae. We have found the eggs of both Smyrna
spp. on Urticastrum mexicanum (leb.) Kuntze, Urera caracasana (Jacquin)
Grisenbach, and U. baccifera (L.) Gaudichaud.

The fast flying females land on the undersurfaces of leaves for ovi-

VoLUME 32, NUMBER 3 167

ie

Figs. 28-31. Smyrna karwinskii: 28, dorsal view of male (note absence of indenta-
tions in anal angles); 29, ventral view of male; 30, dorsal view of female; 31, ventral
view of female.

position. This is in striking contrast to the rest of the Coloburini we have
been able to study, which oviposit mostly on the upper surface of leaves.
Female Smyrna quickly deposit one egg, then move to another plant close
by where the process is repeated. It is not uncommon for the same plant
to be visited again after a period of time. We have observed that females
deposit more than five eggs in sequence before moving away. The plants
used are generally less than 3 m tall.

The rather small eggs are quite hard to locate, due in part to the stinging
properties of the plants, but also to the light green color of the eggs, which
makes them inconspicuous against the color of the leaves. After 5 days
the larva hatches and consumes most of the eggshell, leaving traces of the
wall. It proceeds to feed on the undersurface of the leaf, where it con-
structs a resting perch very close to its feeding place, perpendicular to
the surface. This is a notable deviation from the usual method of the
other perch-making species we know, which always move to the edge of a
leaf to feed, and which construct their resting perch by prolongation of a
vein. In Smyrna spp., second instar larvae do eventually construct the

168 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

perch at the edge of the leaf. From then on, the larvae remain on the
undersurface of leaves until pupation. They pupate on the same plant,
either under leaves or on petioles, after hanging by the anal prolegs for
one day with the thoracic segments incurved ventrally. The pupa hangs
loosely from the pointed cremaster, swinging freely with the faintest
movement of the plant. The colors of the wings of the pharate adult do
not show through the wingcase of the pupal skin when the adult insect
is ready to emerge, due to the dark color of the pupa. After emergence, a
rusty-colored meconium is ejected while the insect expands its wings. The
imago takes a long period of time before its first flight.

Both species show marked sexual dimorphism. At the same time, the
two species are very similar to one another dorsally. The principal differ-
ence is that S. karwinskii has a rounded anal angle on the hindwing,
where S. blomfildia has a small toothed projection.

We have never seen imagos of either species visiting flowers, but we
have seen them feeding on sap flowing from tree wounds, fermenting
fruits, mud puddles, etc., where they spend long periods of time, with
their wings folded dorsally.

Smyrna karwinskii is noteworthy for its seasonal gregarious roosts in
high mountains during the dry season (Muyshondt & Muyshondt, Jr.,
1974). This phenomenon also occurs in Mexico (R. Wind, Chiapas, pers.
comm.; Beutelspacher, 1975). We have never found eggs or larvae of
S. karwinskii in the high mountains during the dry or the rainy season,
even where the foodplants occur locally. It seems that they move down
to lower levels to breed, usually in close proximity to S. blomfildia. The
latter very seldom is found at altitudes over 1600 m and does not have the
communal roosting behavior of S. karwinskii. It is not uncommon to
collect eggs and larvae of both species on the same plant.

The adults of both species behave similarly: they have the same fast
rustling flight, they perch on tree trunks with their heads pointing down,
and the males have a strong territorial defensive attitude, chasing in-
truding butterflies of the same or different species.

To date we have not found cases of parasitism in these species, but we
have very often witnessed predation by Hemiptera (Reduviidae, mostly)
which impale the larvae, leaving only the sagging skin. Another cause of
severe larval mortality in the fields is a disease causing the larva’s body
to burst, releasing a foul-smelling dark fluid.

The foodplants, known locally under one vernacular name common to
the three species, “Chichicaste,” are plants often used as hedges around
coffee plantations, because of the severe stinging caused by the leaves
which deters trespassers.

Urera baccifera grows to a height of 7 m when left alone. The trunks

VOLUME 32, NUMBER 3 169

and older branches are covered with short, wide spines, the younger
branches and the leaves with stinging hairs. The leaves are large, coarse,
round-cordate and roughly dentate. The small greenish flowers grow in
cymes, producing small, translucent, globose fruits with a dark seed
inside. These fruits are much sought after by farm children, who pick
them by beating the shrub with a stick while catching the rain of falling
fruits with a “sombrero.” Thus, hundreds are collected before disposing
of them in situ, much to their pleasure. The juicy fruits have a sweetish-
refreshing taste and alleviate thirst readily.

Urera caracasana grows to about 4 m and is also used in fences. The
leaves are smaller, and of variable shapes; more or less elongate, cordate
at the base and acute at the apex, with close dentation at the edges. The
fruits are red when mature. Both species are used in popular medicine
against venereal diseases. Urticastrum mexicanum is a shrub up to 4 m
tall, with ovate, crenate leaves. The fruits are achenes. All of these
plants have caused painful accidents to tourists unaware of the severe
stinging properties of the otherwise handsome leaves.

DISCUSSION

Smyrna blomfildia is the type-species of the genus Smyrna Hubner,
based on the butterfly originally named Papilio blomfildia by Fabricius
in 1781 (Hemming, 1967). As he very often did, Htibner misspelled the
specific name as “blomfildii,” and this erroneous spelling was subsequently
used by several authors, among them Herrich-Schaffer (1864) and Miller
(1866), with an additional error: “blomfieldii.” Other authors named the
species S. bella Godart and S. pluto Westwood (Seitz, 1921).

The only other reports of the early-stages of this genus that we are
aware of are by Miiller, who gave a short description of a probable 4th
instar larva preserved in alcohol, some rough descriptions based on that
of Miiller’s (Seitz, 1921; Hayward, 1964), and a vague comparison be-
tween the larvae of Colobura and Smyrna by Brown & Heineman (1972).
We believe that ours is the first complete description of the early stages,
with photographic illustrations of both Smyrna blomfildia and karwinskii.

The genus Smyrna has been placed in various unrelated groups based
mostly on the external characters of the perfect insects by many early
authors. Doubleday, Westwood and Hewitson (1849) and Boisduval
(1870), placed it close to Agrias. Herrich-Schaffer (1864) put this genus
in his “familie XI,” together with the related genera Gynaecia and
Callizona, and many unrelated ones, among them Euptoieta, Eunica,
Pyrrhogyra, Ageronia, Peridromia, Amphichlora, etc. Schatz & Rober
(1892), placed Smyrna in their “Gynaecia-Gruppe,” as part of their

170 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

larger “Eunice-Gruppe,” together with Callizona and Gynaecia (Colo-
bura), probably following Herrich-Schaffer.

Today, many authors follow Seitz (1921), who was probably influenced
by Reuter (1898) in placing Smyrna in his “Grupe Gynaeciidae,” together
with Historis, Coea, Pycina, Megistanis, Gynaecia and Callizona, as an
intermediate group between his “Gruppe Epicaliidi” including many
genera (Catonephele, Epiphile, Temenis, Pseudonica, Pyrrhogyra, etc.)
covered by one of us (Muyshondt, 1973a, b, c, d; 1974a) in previous
papers and his “gruppe Hypolimnadidi,” with Hypolimnas misippus L.
Other modern authors include this genus and related genera ( Historis,
Coea, Colobura, etc.) in the Limenitidini which we believe is erroneous.

Different opinions arise, no doubt because of superficial similarities of
the adults, which could very well be due to convergent evolution rather
than close relationship. Examples are well known of convergence in color,
pattern, and shape between unrelated species actually belonging to
different families: i.e. Danaidae, Ithomiidae, Heliconiidae and Pieridae.
These often form Miillerian and Batesian mimicry complexes, as pointed
out by many biologists (Brower, 1972; Brown & Benson, 1974).

The very poor knowledge of the immatures of most tropical butterflies
has led to errors in the association of species. Descriptions of the im-
mature stages are necessary for a more accurate systematic arrangement
of the neotropical Lepidoptera.

Comparison of the eggs of the two species of Smyrna indicates that
they are very closely related. Larvae and pupae are also very similar,
although the larvae of S. karwinskii lack the dorsal row of scoli present in
S. blomfildia. In contrast, the eggs, larvae and pupae of Smyrna differ
considerably from those of Colubura dirce. The immature stages of
Historis odius and Coea acheronta resemble each other closely, but have
nothing in common with Colobura, and only the larval head shape
resembles Smyrna.

There are so many drastic differences between the characteristics of the
early stages of the species of Coloburini studied by us [Colobura dirce L.
(Muyshondt, Jr. & Muyshondt, 1976), Historis odius (Fab.) and Coea
acheronta (Fab.), (Ms. in prep.) ], that we question the correctness of
the taxonomy of the group.

However, we feel that although Smyrna is an aberrant genus in the
Coloburini, it represents a link between the Nymphalini and the other
genera now included in the Coloburini.

By the same token it is also evident that none of these species can be
placed in the Charaxinae, as was done by Boisduval (1870), who placed
Smyrna between Agrias and Prepona, and said that the larvae of

VoLUME 32, NUMBER 3 fal

Aganisthos (=Historis) and Prepona, “sont tout-a-fait semblables” (ex-
actly alike). This is absolutely incorrect! Prepona and Archeoprepona
do resemble each other in the shape of the eggs, larvae and pupae
(Muyshondt, 1973e; Muyshondt, 1976), but neither stage resembles even
remotely the early-stages of Historis (Ms in prep.). To include Smyrna
with the Limenitidini (in which Adelpha belongs) as most modern
authors do, is also incorrect, as they have nothing in common with the
Coloburini during their early stages. With Limenitidini there are certain
imaginal resemblances, but these are not strong enough to place them
together.

It is noteworthy that the larvae of both species of Smyrna construct a
resting perch with frass pellets. Other larvae which use this defensive
strategy construct their perch at the edge of the leaf on which they live:
some of them pile a barrier of excreta mixed with pieces of dry leaf tissue
at the base of the perch (Adelpha spp.); some fasten leaf cuttings with
silk which hang from the perch ( Zaretis, Prepona, Archeoprepona); many
others leave the perch bare (Biblis, Mestra, Catonephele, Epiphile, Nica,
Temenis, Pyrrhogyra, Diaethria, Catagramma, Cyclogramma, Hamadryas,
Colobura, Historis, Coea, Apatura, Marpesia). As far as we have been
able to ascertain only the two species of Smyrna construct a perch on the
underside of a leaf, very close to where the eggshell was consumed.
During the 2nd instar, a new perch is sometimes made at the edge of the
leaf; the other species mentioned also do this. We interpret this behavior
to result from protection afforded to the small larvae by the strong
urticating properties of the foodplants, a factor which by itself might deter
at least some predators. After the 2nd instar the larvae abandon their
perch and wander about the plant on the underside of the leaves. Perhaps
the urticating properties of the plant afford continued protection. It is
to be noted that the profusion of spines displayed by the larvae of Smyrna
spp. from the 3rd instar on do not have urticating properties. Even the
ventral prothoracic gland (adenosma), is not readily extruded as in
Colobura dirce and other species provided with this apparent means of
defense. Thus, it seems that the larvae of Smyrna rely on the protection
granted by the plant itself, rather than on the protection they could derive
from their own spines and odoriferous gland.

One thing puzzles us: although the adults of the two species of Smyrna
are strikingly alike, why is it that the larvae and even pupae of the two
species show important differences, such as the unequal number of rows
of scolii in the larvae, and the different shape of the pupae?

We have seen various larvae and pupae of species belonging to the
same genus, the adults having a common shape but with very disparate

172 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

—

coloration, such as Siproeta stelenes (Young & Muyshondt, 1973) and
V. epaphus (Young, 1972), Heliconius petiveranus and H. charitonius;
Anartia fatima and A. jatrophae. Still others show differences not only
in color, but in the shape of the wings of the adult, as do Catonephele
numilia and C. nyctimus. Yet the larvae and pupae, except for minor
discrepancies, if any, have the same characteristics, indicating that they
undoubtedly both belong to the same genus. We have seen, on the other
hand, species placed in the same genus which have very basic differences
during their early stages, for example Hamadryas februa, H. guatemalena
and H. amphinome; and Anaea eurypile, A. morvus and A. pithyusa,
suggesting that they may belong to different but related genera of one sub-
family or family (Muyshondt & Muyshondt, Jr., 1975a, b, c; Muyshondt,
1974b, 1975a, b). For these reasons, regardless of the striking re-
semblances in adult coloration and shape between Smyrna blomfildia
and S. karwinskii, we suggest they might belong to different genera.
They would then form another case of evolutionary convergence, perhaps
of Millerian mimicry. While S. blomfildia seems to be in the process of
finding its optimum larval characteristics, as suggested by the strking
polymorphism in larval coloration, S. karwinskii apparently has already
achieved stability as it has only one morph. We consider S. karwinskii as
the most evolved of the two, because of the uniformity of characteristics
maintained during its whole life cycle, and thence the model of the two.
S. blomfildia we consider to be the youngest, still an evolving species.
This evolutionary phase seems to have momentarily given S. blomfildia
an advantageous flexibility to overcome adverse conditions which are
reflected in a more abundant population than its more stable relative, S.
karwinskii, at least under the conditions in E] Salvador where the habitats
are continuously and severely affected by human influences, due to the
high population density.

It would be interesting to read an explanation of the present phenom-
enon, where two species evolve divergently during their early stages, yet
seem to evolve advergently during their adult stage. Most of the work
of which we are aware on the evolution of butterflies has concentrated on
their adult stage, disregarding almost completely their early stages, which
perhaps would throw new light on the problem.

ACKNOWLEDGMENTS

We wish to express our gratitude to all the people who helped us with
their guidance, provided reference material, and revised our manuscript
to make it presentable, especially Drs. A. B. Klots, A. B. H. Rydon, J. G.
Sternburg, and G. L. Godfrey. This paper would not have been possible

VoLUME 32, NUMBER 3 Nike

without the great help received from Marilyn and Pierre Muyshondt, who
did much of the fieldwork and observations.

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ScHatz, E. & J. Roper. 1892. Die Familien und Gattungen der Tagfalter, in
Staudinger, O. & E. Schatz, Exotische Schmetterlinge, 2.

Seitz, A. 1921. Macrolepidoptera of the World. Vol. 5. Stuttgart.

STANDLEY, P.C. 1922. Trees and shrubs of Mexico (Fagaceae-Fabaceae). Contrib.
U.S. Natl. Herb. 23, Part 2. 296 p.

Younc, A.M. 1972. The ecology and ethology of the tropical nymphaline butterfly,
Victorina epaphus. I. Life cycle and natural history. J. Lepid. Soc. 26: 155-170.

Younc, A. M. & A. MuysHonpr. 1973. Ecological studies of the butterfly Victorina
stelenes (Lepidoptera-Nymphalidae) in Costa Rica and EI Salvador. Stud.
Neotrop. Fauna 8: 155-176.

Journal of the Lepidopterists’ Society
32(3), 1978, 175-177

SCELIODES LAISALIS (PYRALIDAE): DESCRIPTION OF THE
MATURE LARVA AND NOTE ON ITS FEEDING HABIT

E. O. OGUNWOLU

Entomology Division, National Cereals Research Institute,
P.M.B. 5042, Ibadan, Nigeria

ABSTRACT. Mature laboratory-reared and field-collected larvae of Sceliodes
laisalis Wlk. (Pyralidae) are examined and described and the chaetotaxy illustrated.
A brief note on the feeding habit of this species, an economic pest of Solanum fruits
in southwestern Nigeria, is given.

Taxonomy of immature insects has long been neglected in Nigeria;
mostly, general descriptions of immature stages are contained in publica-
tions on the biology of some species. Where a complex of species is
involved, most taxonomic studies and identifications were of adults. In
contributing to the growth of taxonomy of immatures, I here describe
the larva of Sceliodes laisalis Wlk. and illustrate its chaetotaxy. Its feeding
habit is also described.

Sceliodes laisalis, widely distributed in southwestern Nigeria (Akinlo-
sotu, 1977), is an economic pest of the garden egg fruit, Solanum macro-
carpon L. and S. melongena L. (Solanaceae). Larvae reduce the quality
and quantity of the fruits and the seeds.

My descriptions are based on 15 field collected specimens and 25 larvae
reared in the laboratory. In the following description, names of setae
follow Hinton (1946). To describe the feeding habit at Ibadan, I ex-
amined fruits of ten S. macrocarpon and S. melongena cultivars and those

of Capsicum annuum L.., C. frutescens L., and Lycopersicum esculentum
Mill. for Sceliodes attack.

Description of Larva

General. Length 16.0-22.4 mm, mean = 17.6 mm; width 3.0-4.1 mm, mean =
3.3 mm. Head yellowish brown often with dark brown maculation; ocellar area deep
brown, a fuscous band occurs from post-occiput narrowing towards but not reaching
the ocellar area (Fig. 1B). Prothoracic shield pale with brown spots and deep
brown areas postero-medially; mesothorax suffused with brown; thoracic legs brown.
Abdomen pink dorsally excepting the deep brown anal shield; venter pale. Setal
pinacula light brown. Spiracles circular, yellowish with brown rim; prothoracic and
8th abdominal spiracles each larger than the others.

Head (Fig. 1A). Width 1.1-1.5 mm, mean = 1.2 mm; subspherical in frontal
view. Adfrontal suture reaching the acutely angled vertex. Labrum shallow and
emarginate, mandible with 5 teeth, all but the distal one pointed (Fig. 5), mesal
surface with 4 ridges.

Each of the first, second and the sixth ocellus larger than the others; the second
ocellus closer to the first than to the third. Posterior seta P2 about % the length of P1;

176 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

SDI

—————————

Figs. 1-5. Chaetotaxy of Sceliodes laisalis larva. 1, head, frontal view (A) and
lateral view (B); 2, thoracic segments; 3, first, sixth through ninth abdominal seg-
ments; 4, anal shield; 5, right mandible.

this inserted about mid-length of the head. Seta P1 closer to adfrontal seta AF1 than to
AF2. Distance from AF2—AFa puncture one-third to one-half that from AF2-AF1.
fa punctures well below frontal setae F1; distance between these one-half to three-
fourths that between clypeal setae C2. A2, A3, and LI setae obtuse angled. Vertical
setae V1 slightly antero-ventrad from V2; this closer to V3 than to V1. Ocellar seta Ol
close to the 3rd ocellus, O2 seta ventrad from the Ist ocellus. Distance from genal seta
G1-Ga puncture less than 4% that between Ol—Oa puncture.

VOLUME 32, NUMBER 3 AT

Thorax (Fig. 2). Seta MxD1 outside prothoracic shield. On this seta XD2 dorsad
from and closer to SD1 than to XD1. On meso- and metathorax each of dorsal and
subdorsal setae on the same pinaculum; lateral seta L3 postero-dorsad from LI;
distance from MV2—MV3 about one-third that from MV3-V1. On mesothorax MSDI1
and MSD2 on the same pinaculum but on separate pinacula on metathorax. Pro-
thoracic coxae close but not touching, mesothoracic coxae about the coxal width apart,
and the metathoracic coxae generally more than the coxal width apart.

Abdomen (Fig. 3). D1 and D2 pinacula equal in size and closer on the 8th than
other segments. On the 9th segment, dorsal setae D2 on the same pinaculum; D1, a
thin seta, is slightly closer to SD1 or equidistant between D2 and SD1. Seta MDI
antero-ventrad from D1 on segments 1-8; lateral pinaculum larger on the 8th than on
the other segments.

Subventral setae. 1: 3: 2: 1:1 on segments 1, 2, 7-9, respectively. Distance
between V1 pinacula on the 7th segment more than 2 that of the 9th segment.

Anal shield (Fig. 4). Rounded posteriorly, SD1 setae slightly to well above D1
setae. Prolegs on segments 3-6 with crochets, in biordinal mesoseries, respectively
numbering 16-22, mean = 19; 17-22, mean = 20; 17-23, mean = 22; and 18-24,
mean = 22; and on anal proleg, 14-17, mean = 16.

Description of Feeding Habit

In feeding, larvae tunnel within the fruit which shows no external
sign of damage while larvae are small. The mature larvae exit through
holes made on the fruit. These holes serve as entry sites for decay
organisms, whose activities eventually cause fruit rot. The cultivars of
S. macrocarpon and S. melongena sampled (n = 5 for each) were all
attacked by Sceliodes larvae. Two to five larvae were found within each
fruit. I found no Sceliodes larvae in fruits of Capsicum annuum, C.
frutescens, and Lycopersicum esculentum and neither did Akinlosotu
(1977), but Davis (1964) recorded these as hosts of Sceliodes cordalis
(Dbld.) larvae.

LITERATURE CITED

AxINLosotu, T. A. 1977. A check list of insects associated with local vegetables
in Southwestern Nigeria. Univ. Ife. Inst. Agr. Res. Bull. 8 (In press).

Davis, J. J. 1964. The egg fruit caterpillar. Queensld. Agr. J. 90: 76-78.

Hinton, H. E. 1946. On the homology and nomenclature of the setae of lepidop-
terous larvae, with some notes on the phylogeny of the Lepidoptera. Trans. Roy.
Entomol. Soc. 97: 1-37.

Journal of the Lepidopterists’ Society
32(3), 1978, 178-190

MIGRATION AND RE-MIGRATION OF BUTTERFLIES
THROUGH NORTH PENINSULAR FLORIDA:
QUANTIFICATION WITH MALAISE TRAPS?

THoMAS J. WALKER

Department of Entomology and Nematology,
University of Florida, Gainesville, Florida 32611

ABSTRACT. Malaise traps with single linear barriers perpendicular or parallel to
the axis of the Florida peninsula were operated from 18 Sept. 1975 to 17 Sept. 1976
near Gainesville; insects intercepted by the two surfaces of each barrier were captured
separately allowing them to be scored as flying northward, southward, eastward, or
westward. During the fall, significantly more individuals were caught flying south-
ward than northward for eight species of butterflies: Urbanus proteus (Linnaeus);
Phoebis sennae (Linnaeus); Precis coenia (Hubner); Panoquina ocola (Edwards);
Agraulis vanillae (Linnaeus); Lerema accius (Smith); Urbanus dorantes (Stoll); and
Eurema lisa Boisduval and Le Conte. Estimated net numbers flying southward across
each meter ranged from 3956 (U. proteus) to 33 (E. lisa). During the spring
significantly more individuals were caught flying northward than southward for two
species: P. coenia, A. vanillae. Estimated net numbers were 150 and 10 per m,
respectively. Malaise traps can continuously and effectively monitor insect migration
within the boundary layer.

Long-distance flights by insects are frequent and of theoretical and
practical interest (Williams, 1958; Johnson, 1969; Dingle, 1972). Such
movements are difficult to study because quantification requires identi-
fying flying insects and determining their directions of movement as well
as counting them. Most long-distance flights of insects may occur at
night or at high altitudes making detailed observations impractical, al-
though mass flights above 10 m can be studied with radar (Schaefer,
1976; Riley, 1975). Direct visual observation is useful for large insects
that fly low in daylight. Butterflies have been the most frequent subjects
of such observations (Arbogast, 1966; Baker, 1968b; Balciunas and
Knopf, 1977). Since direct observation is difficult and time consuming,
the resulting data are generally skimpy and likely to be biased by choice
of observation times.

Malaise traps (Southwood, 1966) can complement direct observation
of flights of low flying insects by continuously sampling without the
presence or bias of an observer. Appropriately modified, a Malaise trap
can separate insects flying in one direction when intercepted from those
flying in another direction. I used four such traps to monitor insect
flights within 2 m of the ground for one year near Gainesville, Florida.

‘Florida Agricultural Experiment Station Journal Series No. 457.

VOLUME 32, NUMBER 3 179

Fig. 1 (above). N-—S trap at Green Acres site, looking NNW.
Fig. 2 (below). Heads at one end of trap, showing how insects from the two sides
of the barrier were kept separate.

METHODS

Special Malaise traps (Fig. 1) were constructed and are now com-
mercially available.?, Each trap had a 2.6 X 6 m central barrier. Insects
flew into the trap by either of two 2 X 6 m openings that faced in opposite

2D. A. Focks & Co., P. O. Box 12852, University Station, Gainesville, Fla. 32604.

180 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

directions. Upon striking the central barrier they sometimes worked
their ways toward either end, through a truncated funnel, and into a
receptacle where they were killed by vapors from pieces of dichlorvos-
impregnated plastic.? Traps were made so that insects entering through
one opening remained separated from those entering through the other
opening (Fig. 2). The receptacles were emptied daily or two or three
times per week depending on the numbers caught. Insects caught within
the trap but not present in the receptacles were killed and added to the
appropriate batch.

Mating status of samples of female migrants was determined by dis-
secting for spermatophores.

Traps were set at two sites, 15 km apart: (1) Green Acres Farm,
Agronomy Dept., U. of Fla., 18 Sept—18 Oct. 1975 (4 traps); 19 Oct.—2
Nov. 1975 (2 traps); 10 Apr.—6 June 1976 (2 traps); (2) Archer Road
Laboratory, Entomology and Nematology Dept., U. of Fla. 19 Oct. 1975-
17 Sept. 1976 (2 traps). The first site was an open field with no buildings
or woods within 100 m (Fig. 1). The second was a lawn-like area with
buildings 50 m to the west and east. Traps were set in pairs with one
member of each pair oriented WSW-ENE (perpendicular to the axis of
the Florida peninsula and to the predicted track of migrants—henceforth
called a N-S trap) and the other (an E-W trap), 30 m away, NNW-SSE
(parallel to the axis of the Florida peninsula and at right angles to the
N-S trap).

The insect-catching devices (heads) of the traps were improved during
the first month of the study by changing the receptacles from translucent
polypropylene jars to transparent bags (Fig. 2). Even with improved
heads, a trap captured only a small portion of the insects that flew over
the 6-meter line defined by its barrier. The efficiency of traps was
estimated from counts of individuals captured versus individuals evading
capture during observation periods (4, 5, 12, and 26 Oct., 1975).

RESULTS

For eight species of butterflies the N-S traps caught significantly more
individuals flying southward than flying northward in the fall (Table 1).
For two of the eight, N-S traps caught significantly more individuals
flying northward than flying southward in the spring.

Estimating net numbers moving southward (or northward) each week
or each season requires not only counts of individuals caught but also an
estimate of trapping efficiency. During the four observation periods to
determine efficiency of N-S traps with improved heads, 28 of 314 Urbanus

“e.g. 3 4 6 cm pieces of No Pest or Stable Strip, Shell Chemical Co.

181

VOLUME 32, NUMBER 3

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182 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

proteus (8.9%) were captured. The efficiency for single observation
periods varied from 2 to 12%: 3 of 28 (11%), 19 of 159 (12%), 2 of 88
(2%), 4 of 39 (10%). The lowest efficiency (2%) occurred when the
wind was mainly from the north and the skippers were flying higher
than usual: 70% flew over the trap without first hitting the barrier com-
pared to 38-43% for the other three periods. So few of the other species
were flying that no reliable estimate of capture efficiency was obtained.
During the four observation periods 2 of 8 Precis coenia were captured,
0 of 7 Phoebis sennae, and 0 of 3 Agraulis vanillae. None of these ratios
differ significantly from the 8.9% observed for U. proteus (chi square,
P > 0.05). To simplify calculation of estimates of net displacement
(Table 1 and Fig. 3) while keeping well within the limits suggested by
the data on trapping efficiency, I assumed that each trap with improved
heads captured 10% of the individuals flying over its 6-m line. For
N-S traps with original heads, I assumed a 2.5% efficiency, since when
operated simultaneously with N-S traps with improved heads their catches
were approximately one-fourth as great (e.g. 115 compared to 469 U.
proteus ). Conversion to improved heads was completed 17 Oct. 1975.

E-W traps caught approximately the same number of insects flying
eastward as flying westward. The only significant exception (P < 0.05)
was for U. proteus; 432 were caught in eastward flight versus 177 in west-
ward flight (=2.4:1) during 18 Sept._17 Nov. Such a bias would be ex-
pected if the average track of southbound migrants was east of SSE
(158°), the orientation of the central barrier of E-W traps. Balciunas
and Knopf (1977) determined that the mean track was 147°—i.e., 11°
east of SSE.

Beginning 2 Nov. counts of individuals caught in each end of all traps
were recorded separately. The north and south ends of E—W traps showed
approximately the same biases as the north and south sides of N-S traps.
For example, of the 144 U. proteus trapped 2-17 Nov. in one E-W trap,
137 were caught in the south end. (Data from E-W traps were never used
in estimating net displacement northward or southward.) The fall flights
lasted for six weeks or longer (Fig. 3). The continuous nature of the fall

>

Fig. 3. (Top to bottom) U. proteus, P. sennae, P. coenia, A. vanillae. Weekly
occurrence and net displacement northward or southward 18 Sept. 1975 through 17
Sept. 1976. Downward bars show net displacement southward; upward bars show net
displacement northward. Solid bars indicate a significant (P < 0.05) inequality in
numbers caught flying northward and southward. The lengths of the bars show the
estimated numbers of individuals flying southward (or northward) across 1 m
perpendicular to the axis of the Florida peninsula in excess of those flying in the other
direction. Estimates were made from the numbers caught in 1 or 2 N-S traps. Traps

VoLUME 32, NUMBER 3 183

400, Long-Tailed Skipper (n=2505)

(northward)

(southward)

® go. Cloudiess Sulphur (n=259)
®
z
SS
ea
@
@
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®
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3s 100
Cc
go. Buckeye (n= 302)
ke
za
LJ
=
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O
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)
jeu
w
O
- 3, Gulf Fritillary (n=t3)
LJ
a

SSS = Ga
Fall Winter Spring Summer Fall
1975 ———— 1376 ——— — ____——_—

with original heads (18 Sept.—16 Oct. 1975) were assumed to be 2.5% efficient; traps
with improved heads (11 Oct. 1975-17 Sept. 1976) were assumed to be 10% efficient
(see text). Since the final “week” had 9 days (9-17 Sept.), its estimates were
multiplied by 7/9 prior to plotting. Bars without dots indicate weeks in which numbers
caught flying north and south differed by more than one (N-S traps). Dots indicate
other weeks in which at least one individual was caught (all traps). Bars with dots
(gulf fritillary only) indicate one individual caught flying north or south that week.
Dates for seasons are as in Table 1.

184 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TABLE 2. Sex ratios during fall and spring flights and mating status of females.

Individuals Sex Ratio Females Percent
Species Dates Sexed (M/F ) Dissected Mated
U. proteus 9 Oct.-19 Nov. 926 oe 298 21
P. sennae 9 Oct.-19 Nov. 86 87 37 35
P. coenia 9 Oct.—19 Nov. 116 1.11 AQ, 64
21 Mar.-14 May 94 D207 30 80
P. ocola 9 Oct.-19 Nov. 90 1.00 39 62
A. vanillae 9 Oct.-19 Nov. 53 .66 26 Wi
5 Apr.-l4 May fi 40 ®) 100

1 Significantly different from 1.00 (P < 0.05).
2 Significantly different from 1.00 and from 1.11 (P < 0.05).

flights is not adequately documented in Fig. 3. The small numbers caught
prior to changing to improved heads exaggerated the fluctuations during
the first four weeks. Only for U. proteus were sample sizes large enough
during this period to give acceptable precision to estimates of net dis-
placement. The day-to-day continuity of the southward flights of U.
proteus is revealed by the occurrence of such flights on each of the 52
days that the traps were operated between 18 Sept. and 12 Nov. (Storms
blew down all traps on the other 4 days during this period.) The smallest
one-day catch was 5 (1 flying northward and 4 southward) and the
greatest was 149 (1 and 148 respectively ).

Males and females participated in fall and spring flights. With two
exceptions the sex ratios of migrants did not differ significantly from 1.00
(M/F): Females were significantly in excess of males for U. proteus in
the fall, and males were significantly in excess of females for Precis coenia
in the spring (Table 2). Percent of females that had mated varied with
the season and species from 21 to 100 (Table 2). Mated females generally
contained mature eggs.

Discussion
The Flights

The questions of where trapped individuals of the eight species listed
in Table 1 came from, where they would have gone, and what they would
have done once there cannot be answered from the results of this study
or of earlier studies of one or more of the species (e.g., Williams, 1958;
Arbogast, 1966; Urquhart and Urquhart, 1976; Richman and Edwards,
1976; Correale and Crocker, 1976; Balciunas and Knopf, 1977; and
[sdwards and Richman, 1977). Clues to the answers are provided by the
following information on northern limits and overwintering stages (Howe,
1975; Klots, 1951).

Urbanus proteus (Linneaus) (Pyrginae) occurs northward to Con-

VoLUME 32, NUMBER 3 185

necticut and Arkansas and is not known to overwinter in the U.S. except
in Florida (Howe, 1975). In this study significant southward displace-
ment ceased by late November, yet adults remained abundant about the
flowers of Bidens pilosa Linneaus until a hard freeze occurred 19 Dec.
The next ones seen were three individuals, trapped 3 May, 10 June, and
19 July. Greene (1971) reported that adults were seldom seen at Sanford,
Fla. (140 km SE of Gainesville) from 1 Jan. to 1 July during 1967, 1968,
and 1969.

Precis coenia (Hubner) (Nymphalidae) occurs northward to Wis-
consin, southern Ontario, and New England. Gorlick (in Howe, 1975)
stated that “adults hibernate in winter,” but did not indicate how far
northward such hibernation is known. In this study adults were trapped
during December and February but none was seen or trapped during
January nor for the 104 days between 3 June and 16 Sept. 1976.

Phoebis sennae (Linneaus) (Pieridae) occurs north to Canada but is
not known to overwinter in U.S. except in the Gulf region and Florida.
Adults are seen in Gainesville throughout the winter (though none was
trapped during January 1976; cf. Precis coenis). The two specimens
captured in N-S traps during spring 1976 were flying northward. None
was trapped or seen between 31 Mar. and 28 July.

Panoquina ocola (Edwards) (Hesperiinae) occurs northward to
Arkansas and New Jersey. None was trapped between 16 Dec. and 8
Mar.; 6 were trapped from 8 Mar. through 10 May; none between 10 May
and 3 Sept.

Agraulis vanillae (Linnaeus) (Heliconiidae) occurs northward to
Wisconsin (Schwehr, 1971). No stage survives the winters in Kansas
(Randolph, 1927). After the 19 Dec. 1975 freeze in Gainesville, no adults
were seen or trapped until 12 Feb.

Lerema accius (Smith) (Hesperiinae) occurs northward to New
England and Illinois. During this study the only records were fall catches
(5 Sept.-17 Nov.).

Urbanus dorantes (Stoll) (Pyrginae) is common in southern Texas and
the Greater Antilles but only recently (1969?) became established in
Florida (Knudson, 1974). The Florida population belongs to the Texas
rather than to either of the Antillean subspecies (Miller and Miller,
1970); adults occur all winter in South Florida; in the Gainesville area the
Only records are fall (18 Sept. to 18 Nov.).

Eurema lisa Boisduval and Le Conte (Pieridae) occurs northward to
Quebec and Ontario. Neither adults nor pupae seem to survive the
winters north of 40° (Howe, 1975). When adults appear in Missouri in
late spring, they are “invariably ragged, faded and torn, indicating that

186 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

they may have flown into the area from the south. In the absence of
near freezing temperatures (such as in [south] Florida) there are con-
tinuous broods” (Howe, 1975:372). All records during this study were
in fall (3 Sept—26 Oct.).

The information above is compatible with the hypothesis that each of
the eight species detected moving southward through Gainesville in the
fall breeds farther north than it overwinters. Except for P. coenis and
P. sennae, nothing conflicts with and some data support the more extreme
hypothesis that overwintering occurs no farther north than peninsular
Florida. Either hypothesis requires northward flights in spring or early
summer. The Malaise traps detected such flights only for P. coenia and
A. vanillae. Williams (1958) reported direct observations of northward
spring flights for three of the eight species: P. sennae (Ala.); A. vanillae
(Fla.) and perhaps U. proteus (Fla.). (The other five species are less
conspicuous and/or more difficult to identify on the wing.) The lack
of trapping evidence for northward flights in six of the species might be
attributed to fewer individuals taking part or to different patterns of
flight (e.g. slower and less unidirectional, Nielson, 1960; or above 2 m).

The flights through Gainesville that were documented by Malaise traps
differ significantly from the migratory flights of most insects (Johnson,
1969) and perhaps of most butterflies (Baker, 1968a, 1968b, 1969).

Johnson (1969) emphasized that most long-distance flights by insects
are above the boundary layer—the layer of air near the ground in which
the air movement is less than the insect’s air speed—and cites instances
where long-range dispersal of butterflies may be primarily wind deter-
mined. The thickness of the boundary layer depends on air speed of the
insect, speed of the wind, and degree to which the roughness of terrain or
vegetation slows the air near the ground. The butterflies and skippers
observed during this study and captured in the Malaise traps were gen-
erally, probably always, flying within their boundary layers. When the
wind was blowing in the direction of flight, the migrants flew higher, and
when the wind was blowing counter to the direction of flight, they flew
lower, but butterflies and skippers were never seen flying at wind speeds
greater than their air speeds. When the flights were greatest, the winds
were light and variable. Air speeds for A. vanillae average 18 km/h
(Arbogast, 1966) while those for U. proteus, P. coenia, and P. sennae
average 23, 18, and 20 km/h respectively (Correale and Crocker, 1976;
Balciunas and Knopf, 1977). Flight heights for the same four species
(Over open ground) are generally 0.2-2.0 m (Arbogast, 1966; Edwards
and Richmond, 1977).

Baker (1968c, 1968b, 1969) concluded that at least six and possibly

VoLUME 32, NUMBER 3 187

eight of nine species of British migratory butterflies for which he had data
orient by means of the sun but do not compensate for its movements. In
other words, mean flight direction during the day changes approximately
15°/hr. Such is not the case for the four species that have been studied at
Gainesville. Arbogast (1966) found no significant shift in flight direction
with time of day for A. vanillae nor did Balciunas and Knopf (1977)
for U. proteus. I have similar unpublished observations for P. coenia
and P. sennae. By what means these insects maintain approximately the
same compass direction at all times of day is unknown. Two hypotheses
seem especially worth testing: time-compensated sun orientation (e.g.
Frisch, 1974) and orientation by means of the earth's magnetic field
(e.g. Lindauer, 1977). Malaise-type traps could be used to capture large
numbers of migrants for clock-shifting experiments or for testing with
simulated suns or magnetic fields.

Since the Malaise traps operated continuously and collected small, plain
insects as well as large, showy ones, they had the potential of detecting
migratory flights of species that were rare, inconspicuous, or difficult to
identify on the wing. Four of the eight species detected migrating (Table
1) were such species: P. ocola, L. accius, U. dorantes, and E. lisa. The
methods of trapping and of analyzing the catches could have detected
boundary-layer migratory flights of species in other insect groups—for
example, moths, flies, wasps, and dragonflies. None was detected al-
though low-altitude directional flights of such insects have been observed
elsewhere (Williams, 1958). Species of these groups either did not
migrate through Gainesville at altitudes below 2 m or they migrated in
numbers too small to be detected by one or two 6-m Malaise traps.

The Method

The estimated net displacements northward and southward in Table 1
and Fig. 3 should be evaluated as to precision (repeatability ) and ac-
curacy (correspondence to true value). Different N-S traps at the same
site or at sites 15 km apart gave estimates of net displacement that were
so similar that the traps could not be proved different with the number
of paired observations available (Table 3). The only apparent problem
with precision is variation in trapping efficiency attributable to wind
direction (see above).

Evaluating accuracy depends on comparing the values obtained with
Malaise traps with values obtained by using other methods. Since no
other method of continuous or automatic monitoring has been developed,
the only comparisons that can be made are with direct, visual observations.
The only such observations made were brief and not intended to check

188 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TasLE 3. Comparisons of effectiveness of two N-S traps in detecting net displace-
ment when they were 43 m apart at the same site (Green Acres) and when they were
at two sites 15 km apart.

Net Numbers?

Comparison Number of Trap A Trap B Accept
Species (season) Observations# 6 2s, SID x J ISD2 ree
Same site
U. proteus (fall) 16 15.2 == 20:4 112+114 yes
Different sites
U. proteus (fall) 11 34.9 + 20.4 42.4 + 25.8 yes
P. coenia (spring) 5 Cisse las) 5 Ae ae yes

1 In fall, number intercepted flying southward less number intercepted flying northward; in spring,
the reverse.

2 Standard deviation is used here merely as a measure of variation; distribution of catches was not
assumed to be normal.

3 The null hypothesis (H,) was that traps A and B were sampling the same population with equal
effectiveness. The Wilcoxon Matched-Pairs Signed-Ranks test at P = 0.05 was used (Siegel, 1956).

4 Number of days (U. proteus) or weeks (P. coenia) during which both traps were operative and
at least one individual was caught. For the observations of P. coenia no period shorter than a week
could be used because trap-service dates at the two sites generally coincided but once per week.

the accuracy of trapping estimates; however, the two methods of estima-
tion agree well enough (Table 4) to suggest that the error in trapping
estimates of the four species directly observed is decidedly less than an
order of magnitude. For U. proteus this evaluation has an element of
circularity because direct observation of that species yielded the 10%
trapping efficiency that is incorporated into the formula for estmating net
displacement from trapping results.

Advantages of using Malaise traps to study migration within the
boundary layer include the following: (1) Continual monitoring is prac-
tical. (2) Sensitivity is great enough to detect small-scale flights. (3) Cost
is low enough to permit replication or extensive montoring. (4) Accept-
able precision and accuracy are attainable. (5) Capture of individuals
permits positive identification and determination of sex and mating status.
(6) With modified heads, traps could catch large numbers of individuals
for mark-release studies of destination of migrants or for studies of means
of orientation.

The following are important limitations to using Malaise traps in
studying insect migration (though some can be overcome by modifying
the trap design): (1) Only flights near the ground can be monitored
(modified traps could be suspended from tethered weather balloons).
(2) Traps are damaged by severe storms (hardware cloth or woven wire
could be substituted for the polyester mosquito netting). (3) Information
as to flight direction is crude: + 90° (traps with barriers every 90° or 45°
could be built). (4) Weather factors, such as wind direction, will affect

VOLUME 32, NUMBER 3 189

Taste 4. Comparison of estimates of numbers flying southward across a 1 m
WSW-ENE line, per hour, by direct observation and by trapping at Green Acres site.

Date and Time of Direct Observation

Method of 11 Oct. 1975 12 Oct. 1975 26 Oct. 1975

Species Estimate 1552-1622 1248-1318 1229-1252
U. proteus Direct’ 20.5 OA At 16.0
Trapping” 15.0 23.2 Ul
P. coenia Direct® 1.3 IL 0.9
Trapping? SS) 5.4 0.4
P. sennae Direct® 1.8 0.4 Or
Trapping” 0.7 Zell 0.4
A. vanillae Direct* 0.3 0.4 0.7
Trapping” 0.5 0.5 OL

1 Based on counting individuals crossing a 6 m WSW-—ENE line.

2 Based on the day’s catch of individuals in N—S traps with improved heads. Trapping efficiency
was assumed to be 10% and flight activity was assumed to be spread evenly over 7 hrs: indiv/m/hr
= 10N/w/7, where N is number trapped flying southward and w is width of trap(s) (6 m for 11
and 12 Oct.; 12 m for 26 Oct.).

3 Based on counting individuals crossing a 15 m WSW-ENE line.

height of flight and hence trapping efficiency. (5) An insect entering the
trap from one direction may have a greater probability of beng captured
than one entering from another direction. For example once an insect
has struck the barrier, it may fly toward the brightest light—often the
sun to the south. If the direction of attempted escape is south, an insect
on the south side of the barrier would be more likely to escape than one
on the north side of the barrier. If this bias occurs, it is apparently slight:
For example, Hylephila phyleus (Drury), (Hesperiinae), was caught in
larger numbers than any other nonmigratory skipper or butterfly. Of 198
individuals caught in N-S traps, 111 were captured on the north side and
S87 on the south side. A chi square test reveals no significant bias
CE 005):.

ACKNOWLEDGMENTS

I thank Dana Focks for creating the traps and helping tend them, Dave
Nickle for processing many of the collections, Dale Habeck for help with
techniques, Earl Horner of the Agronomy Department for permitting use
of Green Acres, Tom Emmel for help with identification and literature,
and James Lloyd, James Nation, and Boyce Drummond for criticizing the
manuscript.

LITERATURE CITED

ArsocasT, R. T. 1966. Migration of Agraulis vanillae (Lepidoptera, Nymphalidae )
in Florida. Fla. Entomol. 49: 141-145.

190 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Baker, R. R. 1968a. A possible method of evolution of the migratory habit in
butterflies. Phil. Trans. Roy. Soc. London, Ser. B., Biol. Sci. 253: 309-341.
1968b. Sun orientation during migration in some British butterflies. Proc.

R. Entomol. Soc. London (A) 43: 89-95.

. 1969. The evolution of the migratory habit in butterflies. J. Anim. Ecol.

38: 703-746.

Baucrunas, J., & K. Knorr. 1977. Orientation, flight speeds, and tracks of three
species of migrating butterflies. Fla. Entomol. 60: 37-39.

CorrEALE, S., & R. L. Crocker. 1976. Ground speed of 3 species of migrating
Lepidoptera. Fla. Entomol. 59: 424.

Dincte, H. 1972. Migration strategies of insects. Science 175: 1327-1335.

Epwarps, G. B., & D. B. Ricuman. 1977. Flight heights of migrating butterflies.
Fla. Entomol. 60: 30.

Friscu, K. von. 1974. Decoding the language of the bee. Science 185: 663-668.

GreENE, G. L. 1971. Instar distributions, natural populations, and biology of the
bean leaf roller. Fla. Entomol. 54: 213-219.

Howe, W. H. (ed.). 1975. The Butterflies of North America. Doubleday, Garden
GityvaN.v. G3aup:

Jounson, C. G. 1969. Migration and Dispersal of Insects by Flight. Methuen,
London. 763 p.

Kuors, A. B. 1951. <A Field Guide to the Butterflies of North America, East of the
Great Plains. Houghton Mifflin, Boston. 349 p.

Knupson, E. C. 1974. Urbanus dorantes dorantes Stoll (Hesperiidae): another
example of Florida’s population explosion. J. Lepid. Soc. 28: 246-248.

LinpAvER, M. 1977. Recent advances in the orientation and learning of honeybees.
Proc. XV Int. Congr. Entomol. (Washington, D.C. 1976). pp. 450-460.

Miter, L. D., & J. Y. Mitxier. 1970. Pieris protodice and Urbanus dorantes in
southern Florida. J. Lepid. Soc. 24: 244-247.

NIELSEN, E. T. 1960. A note on stationary nets. Ecology 41: 375-376.

RANDOLPH, V. 1927. Qn the seasonal migrations of Dione vanillae in Kansas. Ann.
Entomol. Soc. Amer. 20: 242-244,

RIicHMAN, D. B., & G. B. Epwarps. 1976. Feeding by four species of migrating
butterflies in northern Florida. Fla. Entomol. 59: 304.

Ritey, J. R. 1975. Collective orientation in night-flying insects. Nature 253:
113-114.

SCHAEFER, G. W. 1976. Radar observations of insect flight. Symp. Roy. Entomol.
Soc. London 7: 157-197.

ScHWEHR, D. W. 1971. Nymphalidae of Wisconsin. J. Lepid. Soc. 25: 139-142.

SIEGEL, SipNEY. 1956. Nonparametric Statistics for the Behavioral Sciences.
McGraw-Hill, New York. 312 p.

SourHwoop, T. R. E. 1966. Ecological Methods with Special Reference to the
Study of Insect Populations. Methuen, London. 391 p.

Urounmart, F, A., & N. R. UrguHarr. 1976. Migration of butterflies along the gulf
coast of northern Florida. J. Lepid. Soc. 30: 59-61.

WituiAMs, C, B. 1958. Insect migration. Collins, London. 235 p.

Journal of the Lepidopterists’ Society
32(3), 1978, 191-197

HYBRIDS BETWEEN CALLOSAMIA AND SAMIA
(SATURNIIDAE)

RICHARD S. PEIGLER!
303 Shannon Drive, Greenville, South Carolina 29615

ABSTRACT. Several crosses between Callosamia promethea and Samia cynthia
were made between 1900 and 1910, and the hybrid specimens are now in museums.
Recently, the cross C. angulifera 6 x< S. cynthia 2 was made, yielding four F; adult
males. Egg hatching rate was high in this cross. The hybrid larvae, cocoons, adults,
and male genitalia all were intermediate between those of the parent species. Other
intergeneric crosses involving these moths produced no adults, but limited information
was obtained from such attempts. The Nearctic Callosamia is closely related to the
Asiatic Samia, perhaps even more closely than to the Nearctic Hyalophora. The chro-
mosome numbers in these genera are all quite different; thus, crosses rarely produce
ova which hatch. Ecological and morphological differences between the genera suggest
that they do not cross in nature.

Between 1900 and 1910 a few lepidopterists succeeded in rearing crosses
between Callosamia promethea (Drury) and Samia cynthia (Drury).
Miss C. G. Soule published her results several times (Soule, 1902, 1906,
and 1907) as did Joutel (1907). Some of the hybrid larvae reared by
Soule were figured by Packard (1914) but apparently Ferguson (1972)
was first to illustrate one of the impressively intermediate adults of such
crosses. Packard (1914) also cited unpublished remarks from Herman
Strecker stating that these hybrids occur in nature. However, considering
that Strecker’s scant descriptions do not coincide with the artificial
hybrids, these remarks should perhaps be discounted. Furthermore, I
indicate later in this paper why such crosses are not likely to occur in
nature.

I have seen hybrids from these early workers in the United States
National Museum and the American Museum of Natural History. Most
of these are obviously true hybrids such as the male of the cross S. cynthia
é X C. promethea ? figured in color by Ferguson (1972). However,
others appear to be pure S. cynthia such as the other “hybrid” which
Ferguson illustrates (a female stated to be of the cross C. promethea 4
x S. cynthia 2). I believe these latter specimens are pure S. cynthia
because it is unlikely that progeny of an intergeneric cross could look so
much like one parent species (in larval, cocoon, and adult traits) and
bear no resemblance to the other. Indeed, Joutel (1907) and Soule (1906)
reveal that they sometimes remated females to males of their own species
in order to initiate oviposition. Future workers who find this practice

1 Museum Associate in Entomology, Natural History Museum of Los Angeles County.

192 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

necessary might try remating females to males which have been artificially
sterilized beforehand. The purpose of this paper is to report results of
similar intergeneric crosses not previously published.

Callosamia angulifera 6 X Samia cynthia

Using hand-pairing and rearing methods which I have described in an
earlier paper (Peigler, 1977), I obtained two fertile matings of this cross
and adults from one of these crosses. The females oviposited freely so that
remating to males of S. cynthia was unnecessary. One of these crosses
employed a male C. angulifera (Walker) from Clemson, South Carolina
and a female S. cynthia from Brooklyn, New York. Of 248 ova laid, only
four larvae eclosed and these died during the first day.

Another male C. angulifera from Clemson was mated to a female S.
cynthia from New Haven, Connecticut. This time the percent hatch was
very high and the larvae were reared on tuliptree ( Liriodendron tulipifera
L.), the food of the father species. Mature larvae were offered ailanthus
( Ailanthus altissima (Mill.) Swingle), the food of the mother species,
and refused it completely. All the larvae were intermediate and exhibited
very little variation, but Soule (1902) reported a “cynthia form” and a
“promethea form” in one of her broods. Although disease killed many
larvae throughout the rearing, 14 cocoons were obtained. Of the 14
cocoons, ten pupated, and ultimately four adult males were secured, the
others dying as pupae. Physiological problems were noted with the
pupal stage. Two individuals remained green contracted larvae (pre-
pupae) in cocoons for over 13 days before finally pupating. Three males
emerged in the autumn, a full year after pupation, and the fourth in the
spring after the second winter. In late May after the second winter, the
two remaining pupae were injected with 12 ug of a-ecdysone in an attempt
to break diapause but they died shortly afterwards.

Descriptions below are based on larvae and pupae preserved in ethyl
alcohol of the hybrids and pure species except the description of the
mature larva which was written with living specimens before me of the
hybrids and mature larvae of S. cynthia and C. angulifera.

First instar. Head like C. angulifera with a light frontal band. Markings on
integument of thoracic and abdominal segments like S. cynthia but weaker. No stripes
as in C. angulifera. Scoli on all segments basically intermediate, tending toward S.
cynthia. Lateral dark patches on all prolegs intermediate.

Second instar. Color deep yellow as in both parent species. Evidence of both the
striping as in C. angulifera and the mottling in S. cynthia. Scoli dark and more like
S. cynthia.

Third instar. Greenish-yellow with darker yellow head and prolegs. Still more like

S. cynthia with mottled integument. Frontal band no longer present, which would be
in pure C, angulifera of the same instar.

VOLUME 32, NuMBER 3 une}

Fig. 1. Mature larva of C. angulifera $ x S. cynthia 2 on tuliptree. (Photo by
G. R. Carner. )

Fig. 2, Adult male of C. angulifera 6 x S. cynthia °. (Photo by E. V. Gage.)

194 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fourth instar. Very similar to the fifth instar but with whitish powder covering
integument.

Fifth instar (Fig. 1). Color lighter than C. angulifera but waxy white powder
minimal or absent. Head like both parents; the heads of C. angulifera and S. cynthia
are much alike in size, color and shape. Yellow on top of prothoracic segment as in
S. cynthia (absent in C. angulifera). Thoracic legs darker green-yellow than in C.
angulifera. Lateral stripe concolorous with body (like S. cynthia) or slightly yellowish
(very yellow in C. angulifera). No black spots on the integument as in S. cynthia but
never in Callosamia. Spiracles light blue. Prolegs mostly like C. angulifera but with
some yellow as in S. cynthia. Anal claspers without blue of S. cynthia; the black ring
of C. angulifera reduced to a posterior black stripe. Anal plate with two lateral black
spots of some C. angulifera but lacking the black streak between. Dorsal scoli (tu-
bercles) of meso- and metathoracic segments orange with yellow-green base, longer
than in C. angulifera. Dorsal scoli on first abdomenal segment orange or yellowish-
orange on tip, higher yellow-green base. All other dorsal scoli yellow, one greenish
larger one on eleventh segment. Subdorsal scoli small, raised, pointed blue tubercles;
in some individuals the ones on third and fourth segments with black at base. Sub-
spiracular tubercles all small and blue with black bases; the one on first segment a
black dot as in C. angulifera. Thoracic segments each with a subventral black button
as in both parent species.

Pupa. Anal angles of forewings lying on fourth abdominal segment as in Callosamia.
The maxillary area between antennal covers much more like S. cynthia (see Mosher,
1916).

Cocoon. About two-thirds with attachments to stem. Intermediate, but variable
shapes tending to S. cynthia and dark brown coloration quite like C. angulifera.

Male (Fig. 2). Antennae and frons like C. angulifera in size and coloration. Scaling
on body intermediate in color. No dorsal white tufts on abdomen but lighter scaling
in first two abdominal tergites as in S. cynthia. Legs without white scaling. Ante-
median line intermediate. Postmedian line with maroon replacing light pink in S.
cynthia and golden suffusion in C. angulifera. Ground color and submarginal orna-
mentation of forewing and hindwing all intermediate. Apical black patch larger like
C. angulifera. Discal mark in each wing more like S. cynthia but with an anterior tooth
suggesting C. angulifera. Overall coloration of underside much closer to C. angulifera.

Male genitalia (Fig. 3). Intermediate in general but more like C. angulifera.
Uncus larger as in S. cynthia, and saccular lobe of valve less produced as in S. cynthia.
Median lobe like S. cynthia but wide costal lobe like C. angulifera, with no evidence of
the indentation which S. cynthia so clearly has in the costal lobe. Anellus more like
S. cynthia but transtilla (gnathos) weaker as in C. angulifera. Aedeagus shorter than
either parent species and intermediate in width. Only one hybrid dissection studied.

Unfortunately, no attempt was made to test fertility of these F, males.

Of the four males obtained, one is in the collection of Dr. Claude Lemaire

of France, another in the Los Angeles County Museum, and two are in
my collection.

Additional Crosses

Limited information is available on other intergeneric crosses and it
seems appropriate to report it here. These include Callosamia securifera
(Maassen) 4 x Samia cynthia 2 and Callosamia promethea 6 Xx Hyalo-
phora cecropia (.) ?. None of these resulted in adult imagines.

Using a male of C. securifera from Berkeley County, South Carolina

VoLUME 32, NUMBER 3 195

Fig. 3. Male genitalia of C. angulifera 6 x S. cynthia 9.

and a female of S. cynthia from Paris, 58 ova were secured. Only six
ova hatched but most other embryos reached full development even
though they failed to hatch. This became apparent when viewing the
batch of ova after they had been stored in ethyl alcohol. The six larvae
were offered tuliptree but were very weak and expired within a few hours.

196 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

They had banded heads and dorsal segmental stripes as in pure C.
securifera, but the scoli were more like S. cynthia.

Mr. Laurence R. Rupert of Sardinia, New York very kindly provided
information on a cross he made between a male C. promethea and a
female H. cecropia over 60 years ago:

“. . After holding the abdomens together for a time, I succeeded in inducing a
mating. The cecropia produced a good number of eggs, but only 8 or 9 hatched.
The young larvae at first resembled those of cecropia, but as time went on they grew
more and more to resemble those of promethea. I fed them lilac [Syringa vulgaris L.]
since I knew that was a favorite food of both parents.

Only two larvae survived to make cocoons, and the two cocoons were very different.
One was quite hard, resembling in shape a very small cecropia cocoon of the hard
non-puffy type. The other was irregular in shape with a strip of silk along a stem,
like a promethea cocoon, but weaker. No moth emerged from either, and I never
bothered to open the cocoons to see whether the larvae pupated.”

The same season during which I succeeded with the C. angulifera 4
x S$. cynthia 2 hybrids, numerous other matings were made which re-
sulted in no eclosion of ova. Adults of Samia cynthia, S. canningii
(Hutton), H. cecropia, and hybrids and all three species of Callosamia
were used in various combinations. In general, the male genitalia of C.
promethea and H. cecropia are so large that females of other species are
often killed by mating with them. The genitalia of other Callosamia males
are very compatible with Samia females and such crosses should be tried
whenever possible. I have been unable to accomplish hand-pairings in
which the male was S. cynthia.

DIscussION

The value of hybridization as a tool in elucidating phylogeny has been
well documented by Hubbs (1967). The fact that Samia is Asiatic and
Callosamia is Nearctic may have caused earlier authors to overlook the
affinities of the two groups. The close relationship based on morphological
studies has been pointed out by Lemaire & Peigler (1978). Additionally,
tuliptree, the main food of Callosamia, is an excellent food for pure
S. cynthia, the larvae growing much faster than they would on other
substitutes for their preferred host, ailanthus (pers. obs.). This becomes
even more significant when one finds that larvae of the genus Hyalophora,
considered closest to Callosamia, fail to survive on tuliptree (Scarbrough
et al., 1974). Intrageneric introgression sometimes proves to be advan-
tageous to populations of organisms, but one can hardly see any possible
advantage in intergeneric hybridization. Therefore, some specialized bio-
chemical isolation mechanism may be in effect for the two sympatric
genera of North America, not shared by the Asiatic Samia.

VoLUME 32, NUMBER 3 197

The chromosome number of C. promethea is 19, that of the species of
Hyalophora is 31, and the various species of Samia is 13 and 14 (Robinson,
1971). Thus the pairing of sex chromosomes and autosomes possibly
becomes critical when hybridizing these related genera, and this may
explain several results such as lack of females (the heterogametic sex in
Lepidoptera) in many hybrid broods, the rarity with which ova hatch
in such crosses, and the different results sometimes obtained with
reciprocal crosses (see Peigler, 1977).

The question arises as to whether such crosses ever occur in nature so
that a collector might encounter such wild hybrids. The larger genitalia
of C. promethea and its diurnal mating behavior argue against the
possibility of this species mating with S. cynthia. The genitalia and mating
times of C. angulifera and S. cynthia are much more compatible, but a
pheromone difference exists wherein females are unlikely to attract males
of the other species. Also, S. cynthia exists almost exclusively in large
cities where C. angulifera populations are quite unlikely to occur with any
regularity, if at all. Even after being in North America over 100 years, S.
cynthia still appears to be predominantly on ailanthus in urban areas.

LITERATURE CITED

Fercuson, D.C. 1972. Bombycoidea, Saturniidae (in part). In R. B. Dominick et
al., The Moths of America North of Mexico, fasc. 20.2B: 155-269, pls. 12-22.

Husss, C. 1967. Analysis of phylogenetic relationships using hybridization tech-
niques. Bull. Natl. Inst. Sci. India 34: 48-59.

JoureL, L. H. 1907. Philosamia cynthia and Callosamia promethia [sic] crosses.
J. N. Y. Entomol. Soc. 15: 101-103.

Lemarre, C. & R. S. PEIGLER. 1978. A study of Samia watsoni (Attacidae). J.
Res. Lepid. In press.

MosHer, E. 1916. The classification of the pupae of the Saturniidae. Ann. Entomol.
Soc. Amer. 9: 136-158.

PacKxarp, A. S. 1914. Monograph of the bombycine moths of North America, part
3 (T. D. A. Cockerell, ed.). Mem. Natl. Acad. Sci. 12: 1-516.

PricLer, R. S. 1977. Hybridization of Callosamia (Saturniidae). J. Lepid. Soc.
31: 23-34.

Roprinson, R. 1971. Lepidoptera Genetics. Pergamon Press, New York. 687 p.

ScARBROUGH, A. G., G. P. WALDBAUER & J. G. STERNBURG. 1974. Feeding and sur-
vival of cecropia (Saturniidae) larvae on various plant species. J. Lepid. Soc.
28: 212-219.

SouLEe, C. G. 1902. Notes on hybrids of Samia cynthia and Attacus promethea.
Psyche 9: 411-413.

1906. Notes on moths. Entomol. News 17: 395-397.

1907. Some experiments with hybrids. Psyche 14: 116-117.

Journal of the Lepidopterists’ Society
32(3), 1978, 198-206

RHOPALOCERA OF WEST VIRGINIA?

BaAsTIAAN M. Drees? AND LINDA BUTLER
Department of Entomology, West Virginia University, Morgantown 26506

ABSTRACT. A current summary of the records of the butterflies (Rhopalocera )
of West Virginia is given. Published records, including references to the state’s fauna
discussed in William Henry Edwards’ Butterflies of North America, provided distribu-
tion data for 71 species. These records are indexed in order that the sources of in-
formation may be easily traced. Extensive collecting by the authors from 1972 to
1977, records of museum specimens and contributions of data by other collectors
resulted in records of 40 additional species, making a total of 111 butterflies now
known from West Virginia. A listing of 34 species which are expected to occur in the
state is also included.

The distribution of Rhopalocera has been of continuous concern to
lepidopterists. Within the past century, many state and national checklists
have been compiled. This is an important step toward monitoring popula-
tion levels and range changes of species. Recently, these lists have become
valuable to conservationists attempting to define endangered species.
Without the basic information these checklists provide, few studies re-
lating to other aspects of Rhopalocera can be undertaken.

While butterflies of most of the states have already been studied, the
fauna of West Virginia has, until now, been neglected. Faunal studies of
the states bordering West Virginia allude to species which should occur
here. With regard to West Virginia, Field et al. (1974) list 38 publica-
tions, 34 of which were written by W. H. Edwards. The information in
the Edwards publications is best summarized in his three volumes of
Butterflies of North America (1868, 1884 and 1897). Edwards mentioned
35 species with specific reference to West Virginia in his voluminous
work. Many of these 35 species were taken at Coalburg in Kanawha
County. Brooks (1905) listed 24 species in the Cranberry River area.
Additional published records of West Virginia butterflies not given by
Field et al. (1974) include those of Holland (1913), Burns (1964), Clench
(1972), Irwin (1972) and those by contributors to the Field Summaries
in the Lepidopterists’ News and the News of Lepidopterists’ Society:
Allen (1976), Boscoe (1976), Butler (1976), Drees (1974, 1976), Jensen
(1976), Lavy (1974), Nicolay (1953), Parshall (1974), Preston (1951)
and Showalter (1975). These publications provide distributional data for
71 species of butterflies which occur in West Virginia. Clark and Clark

. be, Cain laa the approval of the Director of the West Virginia University Experiment Station
’aper No. 1500, rf

* Current address: Department of Entomology, Ohio State University, Columbus.

VoLUME 32, NUMBER 3 199

WEST VIRGINIA

YX Coalburg

Fig. 1. Map of West Virginia with county designations. The site of Coalburg in
Kanawha County was the location of many of the W. H. Edwards state records.

(1951) further noted that “a few early records from ‘Virginia’ refer to
West Virginia which was part of Virginia until 1863.”

Intensive collection throughout the state for the past five years by the
authors and contributions of collection records by T. J. Allen, J. W.
Pamine eae. Cole! C. V: Covell; jr., J. M. Flanery, P. Francis, J. D.
Hacker, R. Lavy, R. H. Lindsey, T. L. Mason, A. H. Showalter, R. E.
Stanford and M. D. Taylor have added to the construction of a more com-
plete statewide list. Data from these sources and data obtained by the
authors from Carnegie Museum of Natural History, National Museum
of Natural History (USNM), West Virginia Department of Agriculture,
and West Virginia University Entomology Department collections have
resulted in 40 previously unpublished state records. Thus, at present, a
total of 111 species is known from West Virginia.

Specimens in the possession of the authors in the West Virginia Uni-
versity insect collection were identified, and the more difficult groups
were confirmed by J. M. Burns, USNM (Hesperiidae); H. K. Clench,
Carnegie Museum of Natural History (Lycaenidae and Nymphalidae);
and W. D. Field, USNM (Lycaenidae). Advice from these sources as well

200 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

as that from H. A. Freeman and A. B. Klots has not only enhanced the
accuracy of this paper, but has made valuable contributions to the listing
of 34 additional species expected to be found in West Virginia. All of
the species in the following list have already been reported from one
or more of the bordering states (Covell, 1967, 1974; Covell and Straley,
1973; Fales, 1974; Clench, unpublished; Wood and Gottschalk, 1942).

Hesperiidae: Lerodea eufala (Edwards), Atrytonopsis hiana hiana (Scudder),
Euphyes dion dion (Edwards), E. dukesi (Lindsey), E. conspicua conspicua (Edwards),
Poanes massasoit massasoit (Scudder), P. viator (Edwards), Polites vibex vibex
(Geyer), Hesperia metea Scudder, Thymelicus lineola (Ochsenheimer ), Lerema accius
(Smith), Erynnis persius persius (Scudder), E. zarucco zarucco (Lucus), and perhaps
Panoquina ocula (Edwards), Calpodes ethlius (Stoll), Poanes yehl (Skinner), Prob-
lema byssus (Edwards), Hesperia attalus attalus (Edwards), Thorybes confusis Bell
and Urbanus proteus (Linnaeus ).

Pieridae: Colias cesonia (Stoll) and Nathalis iole Boisduval.

Riodinidae: Calephelis muticum McAlpine.

Lycaenidae: Satyrium caryaevorus (McDunnough), Euristrymon ontario ontario
(Edwards), and perhaps Satyrium acadica acadica (Edwards), Callophrys irus irus
(Godart), Panthiades m-album (Boisduval and LeConte) and Lycaeides melissa
samuelis Nabokov.

Nymphalidae: Anaea andria andria Scudder, Nymphalis vau-album (Denis and
Schiffermiiller ) and Chlosyne harrisii harrisii (Scudder).

Satyridae: Perhaps Lethe creola (Skinner) and Euptychia areolata areolata
(Smith).

The following list of the Rhopalocera of West Virginia follows basically
the sequence and nomenclature of Dos Passos (1964) and his revisions
(1969, 1970), but includes further changes and additions by Cardé et al.
(1970), Clench (1972, 1975), Field (1971), Freeman (1973) and Howe
(1975). County records and extreme collecting dates for more than 1340
specimens are included. Location of counties is shown on the West
Virginia map (Fig. 1). Index numbers given in parentheses in the species
list refer to notes at the end. Counties with index numbers do not
necessarily represent single collection records. The authors would greatly
appreciate any corrections or additional data for future revision of this
initial summary.

Hesperiidae

Amblyscirtes samoset (Scudder )—Monongalia, Pocahontas (2); V-8.

A. vialis (Edwards )—Mineral, Monongalia; V-4 to 24.

Euphyes bimacula (Grote and Robinson )—Pendleton (11), Webster; VIII-5 to 8.

I. vestris metacomet ( Harris )—Braxton, Pocahontas, Randolph; VI-30 to VII-10.

Poanes massasoit hughi Clark (5).

P. hobomok ( Warris )}—Barbour, Berkeley, Braxton, Brooke, Grant, Mineral, Mon-
ongalia, Pendleton, Pocahontas, Preston, Randolph; V-12 to VII-4.

P. zabulon (Boisduval and LeConte)—Monongalia, Ritchie (5, 12); V-20 to VII-18.

Atrytone delaware delaware (Edwards )—Monongalia, Pendleton (11); VII-22.

Atalopedes campestris (Boisduval )—(5), Calhoun, Putnam; IX-19 to X-11.

Pompeius verna verna (Edwards )—(10) Braxton, Fayette, Gilmer, Hampshire,
Hardy, Mercer, Mineral, Monongalia, Pendleton, Pocahontas; VI-24 to VIII-10.

VoLUME 32, NUMBER 3 201

Wallengrenia egeremet (Scudder)—Berkeley, Braxton, Fayette, Mineral, Mon-
ongalia, Pendleton; VI-8 to VII-11.

Polites coras (Cramer )—Berkeley, Cabell, Grant, Mineral, Monongalia, Pendleton,
Preston, Randolph, Webster, Wetzel; V-20 to IX-9.

P. themistocles (Latreille )—Cabell, Monongalia; V-23 to VIII-13.

P. origenes origenes (Fabricius )—Mineral, Putnam; VI-25 to IX-19.

P. mystic (Scudder )—Preston, Randolph, Webster; VI-5 to VII-8.

Hesperia sassacus sassacus Harris—Preston, Randolph; VI-5 to 19.

H. leonardus Harris—(5), Kanawha, Lewis; VIII-1 to 7.

Hylephia phyleus (Drury )—Kanawha, Lewis; VIII-22.

Ancyloxypha numitor (Fabricius )—Berkeley, Brooke, Gilmer, Grant, Mason,
Monongalia, Preston; V-24 to IX-15.

Nastra lherminier ( Latreille )—Monongalia; VII-3.

Pholisora catullus (Fabricius )—Monongalia; VII-22.

Pyrgus centaureae wyandot (Edwards )—Hardy, Kanawha; IV-16.

P. communis communis (Grote )—Monongalia, Pendleton; VIII-25 to X-2.

Erynnis icelus (Scudder and Burgess)—Barbour, Berkeley, Fayette, Grant,
Hampshire (1), Kanawha, McDowell, Mineral, Monongalia, Pendleton (11),
Preston, Pocahontas (1), Summers, Tucker; IV-11 to VI-25.

E. brizo brizo (Boisduval and LeConte )—Hampshire (1), Kanawha, Monongalia
(1), Pendleton (11); IV-22 to VI-2.

FE. lucilius lucilius (Scudder and Burgess )—(1), Pendleton (11); V-5.

E. baptisiae (Forbes )—Berkeley; VIII-3.

E. martialis (Scudder )—Hampshire (1); V-29.

E. horatius (Scudder and Burgess )—Berkeley, Hampshire (1), Hardy, Upshur
(1); V-7 to VII-3.

E. juvenalis juvenalis (Fabricius )—Barbour, Grant, McDowell, Hampshire (1),
Monongalia (1), Pendleton (11), Pocahontas (1), Preston, Summers, Tucker,
Webster; IV-15 to VIII-3.

Staphylus mazans hayhurstii (Edwards )—(5).

Thorybes bathyllus (Smith)—Mineral, Monongalia, Preston; VI-27 to VII-13.

T. pylades (Scudder )—Gilmer, McDowell, Mineral; V-21 to 24.

Achalarus lyciades (Geyer)—(7), Kanawha, Lewis, Mineral; V-24 to VIII-7.

Autochton cellus (Boisduval and LeConte )—-Kanawha (5); V-15.

Epargyreus clarus clarus (Cramer )—Barbour (15), Berkeley (15), Braxton (15),
Fayette, Gilmer, Grant, Hardy, Kanawha, McDowell, Mercer. Mineral, Mingo,
Monongalia, Pendleton (11), Preston, Wayne; V-7 to VIII-18.

Papilionidae

Battus philenor philenor (Linnaeus )—Barbour (15), Braxton, Boone, Greenbrier.
Hardy, Marion, McDowell, Randolph, Summers, Wayne, Webster (3); IV-30
to X-4.

Papilio polyxenes asterius Stoll—Braxton, Fayette (15), Greenbrier, Kanawha,
Lincoln, Monongalia, Pocahontas, Randolph; V-11 to IX-13.

P. cresphontes cresphontes Cramer—Jefferson (15), Monongalia; VII-3 to IX-1.

P. glaucus glaucus Linnaeus—Barbour, Berkeley, Boone, Braxton, Brooke, Clay,
Fayette, Gilmer, Grant (15), Hardy, Kanawha (5), Marion, McDowell (15),
Mercer, Mineral, Mingo, Monongalia (12), Preston, Putnam, Randolph, Sum-
mers (15), Tucker, Upshur, Wayne, Wetzel; IV-16 to IX-19.

P. troilus troilus Linnaeus—Barbour (15), Berkeley, Braxton, Boone, Cabell,
Fayette, Gilmer (15), Grant (15), Hardy, Hampshire, Kanawha, Marion,
McDowell (15), Mercer, Mineral, Mingo, Monongalia (12), Pocahontas,
Preston, Randolph, Summers (15), Taylor, Wayne, Webster (3), Wetzel,
Wood; IV-15 to IX-20.

Graphium marcellus (Cramer )—Boone, Braxton, Clay, Gilmer, Hampshire, Jack-

©
bo

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

son, Kanawha (5), Monongalia, Randolph, Ritchie, Roane, Wayne, Webster
(3); IV-21 to VITII-22.
Pieridae

Pieris protodice protodice Boisduval and LeConte—Kanawha (5), Webster (3);
VII-2 to VIII.

P. virginiensis Edwards—Kanawha (5), Monongalia, Preston, Randolph (14),
Tucker, Webster; IV-10 to V-14.

P. rapae (Linnaeus )—Berkeley, Braxton (15), Cabell, Grant, Hampshire, Mingo,
Mineral, Monongalia, Preston, Randolph, Gilmer, Roane, Webster (3); III-19
to IX-12.

Colias eurytheme eurytheme Boisduval—Barbour, Berkeley, Braxton, Gilmer
(15), Mercer, Mingo, Mineral, Monongalia, Pendleton, Pocahontas, Preston,
Putnam, Randoloh, Summers (15), Tucker, Wetzel; IV-27 to X-25.

C. philodice philodice Godart—Cabell, Braxton, Gilmer, Grant, Greenbrier, Hardy,
Kanawha (5), Marion, Mason, McDowell, Mercer, Mingo, Mineral, Monongalia,
Pocahontas, Preston, Putnam, Randolph, Summers (15), Tucker, Webster (3),
Wetzel; V-20 to X-25.

C. interior interior Scudder—Randolph (13); V-7 to VII-31.

Phoebis sennae eubule (Linnaeus )—Upshur; VI-12.

Eurema lisa Boisduval and LeConte—Boone, Clay, Kanawha, Lewis, Monongalia;
VIII-6 to X-9.

E. nicippe (Cramer)—Hardy, Kanawha, Lewis, Pendleton, Summers; VIII-6
to X-4.

Anthocaris midea Hiibner—Greenbrier, Hampshire (12), Hardy, Kanawha (5),
Mineral, Monongalia, Pendleton, Wood, Pocahontas; III-30 to V-30.

Euchloe olympia olympia (Edwards )—Hampshire (12), Hardy, Kanawha (5),
Mineral; IV-16 to V-10.

Riodinidae

Calephelis virginiensts virginiensis (Guérin-Méneville )—(5).

C. borealis (Grote and Robinson )—Greenbrier (7), Hampshire, Monongalia; V-22
to VII-14.

Lycaenidae

Harkenclenus titus mopsus (Hiibner )—Fayette, Mineral, Pendleton (11), Poca-
hontas, Randolph; VI-25 to VII-11.

Satyrium liparops strigosa (Harris )—Fayette, Kanawha (5), Monongalia; VI-11
to VII-7.

S. calanus falacer (Godart )—Fayette, Mineral, Monongalia, Pendleton (10),
Pocahontas; VI-25 to VII-19.

S. edwardsii (Saunders )—Greenbrier (14), Hampshire, Randolph; VII-12 to 26.

Calycopis cecrops (Fabricius )—Lewis, Nicholas; VII-8 to 14.

Callophrys polios polios Cook and Watson—Kanawha; IV-20.

C. henrici henrici (Grote and Robinson )—Hampshire, Kanawha; IV-25 to VII-28.

C. augustinus croesioides Scadder—Hampshire, Kanawha; IV-23 to V-3.

C. niphon niphon (Hibner)—Berkeley, Hampshire (12), Pendleton (11); V-5
to 28.

C. gryneus gryneus (Hibner )—Hampshire; IV-25 to V-10.

Strymon melinus humili (Harris )—Berkeley, Fayette, Greenbrier (14), Hardy,
Mineral, Monongalia, Pendleton, Pocahontas, Randolph, Wood; VI-8 to IX-9.

Erora laeta (Edwards )—Cabell (6), Greenbrier (6), Kanawha (5, 6), Pendleton
(11), Randolph; V-5 to VII-26.

Feniseca tarquinius tarquinius (Fabricius )—Mineral, Monongalia, Pendleton
(11), Ritchie (12), Upshur, Webster (3): V-4 to IX-11.

Lycaena hyllus (Cramer )—Brooke, Monongalia; V-28 to IX-13.

L. phlaeas americana Harris—Barbour, Hampshire (12), Marion, Mason, Mineral,

VOLUME 32, NUMBER 3 203

Monongalia, Pendleton (11), Randolph, Taylor, Webster (3), Wood; V-11
to IX-30.

Everes comyntas comyntas (Godart)—Braxton, Grant, Greenbrier, Hardy, Min-
eral, Monongalia, Pocahontas, Preston, Randolph, Webster (3), Wetzel, Wood;
V-2 to X-4.

Glaucopsyche lygdamus nittanyensis Chermock—Greenbrier, Kanawha (5), Ohio,
Pendleton (11), Webster, Wood; V-5 to VI-15.

Celastrina argiolus pseudargiolus (Boisduval and LeConte)—Braxton, Boone,
Cabell, Fayette, Gilmer, Grant, Greenbrier, Kanawha (5), McDowell, Mineral,
Monongalia, Pendleton (11), Preston, Tucker, Wayne, Webster (3) Wetzel;
III-30 to VIII-20.

C. ebenina Clench—Kanawha (4,5), Pendleton (4); IV-17 to V-5.

Libytheidae

Libytheana bachmanii bachmanii ( Kirtland )—Kanawha (5), Mercer (14), Mon-

ongalia, Randolph; VII-26 to IX-10.
Nymphalidae

Asterocampa celtis celtis (Boisduval and LeConte )—Kanawha, Mercer, Mineral
(14), Monongalia, Morgan, Wayne, Wood (15); V-24 to IX-10.

A. clyton clyton (Boisduval and LeConte )—Mingo, Monongalia, Wayne; VII-17
tor27,

Limenitis arthemis arthemis (Drury )—Mineral (15).

L. arthemis astyanax (Fabricius )—Berkeley, Boone, Braxton (15), Gilmer (15),
Grant, Greenbrier, Hardy, Marion, Mason, McDowell, Mercer, Mineral, Mingo,
Monongalia, Pocahontas, Preston, Wayne, Webster (3), Wetzel; V-12 to X-9.

L. archippus archippus (Cramer )—Barbour (15), Boone, Braxton, Grant, Kana-
wha (5), Mason, McDowell (15), Mineral, Mingo, Monongalia, Randolph,
Upshur, Webster (3), Wetzel, Wood; V-12 to IX-13.

Vanessa atalanta (Linnaeus )—Braxton (15), Hampshire (15), Marion, Monon-
galia, Pocahontas, Preston, Randolph, Tucker (15), Webster (3), Wetzel; V-1
to IX-20.

Cynthia virginiensis (Drury )—Barbour, Grant (15), Hampshire, Kanawha, Mc-
Dowell, Mercer, Monongalia, Randolph, Tucker, Webster (3), Wood; V-7 to
VITI-23.

C. cardui (Linnaeus )—Kanawha, Monongalia, Randolph; VII to VIII-30.

Junonia coenia ( Hiibner )—Grant, Randolph; IX.

Nymphalis milberti milberti (Godart )—Monongalia; X-21.

N. antiopa antiopa (Linnaeus )—Brooke (14), Greenbrier, Kanawha, Mineral,
Monongalia, Pocahontas, Randolph, Tucker (15), Webster (3); III-10 to IX-4.

Polygonia interrogationis (Fabricius)—Barbour (15), Boone, Braxton, Gilmer
(15), Grant, Kanawha (5), Marion, Mineral, Mingo, Randolph, Ritchie,
Webster, Wetzel: IV-18 to IX-20.

Polygonia comma (Harris)—Barbour (15), Boone, Kanawha (5), Marion,
McDowell (15), Monongalia, Monroe, Pendleton (11), Putnam, Randolph,
Ritchie, Webster (3); V-5 to IX-20.

P. faunus faunus (Edwards )—Fayette (5), Webster (3); VIII.

P. progne (Cramer )—Cabell, Mercer, Mineral, Mingo, Monongalia, Pendleton,
Wayne, Webster; VI-29 to X-14.

Chlosyne nycteis nycteis (Doubleday )—Boone, Braxton, Fayette, Grant, Hamp-
shire, Hardy, McDowell, Mercer, Mineral (14), Monongalia, Pendleton, Ritchie,
Wetzel; VI-18 to VIII-28.

C. gorgone carlota ( Reakirt )—Kanawha; V-4.

Phyciodes tharos tharos (Drury )—Barbour, Berkeley, Boone, Cabell, Fayette,
Greenbrier, Hampshire, Hardy, Kanawha (5), Marion, McDowell, Mineral,
Mingo, Monongalia, Pocahontas, Preston, Putnam, Randolph, Summers, Wayne,
Webster (3); IV-27 to IX-19.

204 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

P. batesii (Reakirt )—Kanawha; V-22 to 28.

Euphydryas phaeton (Drury)—Kanawha (5), Monongalia, Randolph (14);
VI-1 to VII.

Boloria bellona bellona (Fabricius )—(10), Boone, Braxton, Greenbrier, Hamp-
shire, Hardy, Jackson, Kanawha, Marion, Mineral, Monongalia, Pendleton (11),
Pocahontas, Randolph, Webster (3), Wood; IV-18 to X-4.

B. selene myrina (Cramer )—Grant, Kanawha, Randolph; V-24 to VII.

Speyeria idalia (Drury )—Braxton, Clay, Hampshire, Lewis, Monongalia, Pen-
dleton (11); VI-25 to IX-20.

S. atlantis atlantis (Edwards )—Monongalia, Pendleton (11), Randolph, Webster
(3); VI-18 to IX-14.

S. diana (Cramer )—Boone (14), Fayette (5), Kanawha (5), Mercer, Webster
(3); VI-13 to VII-2.

S. cybele cybele (Fabricius )—(10), Berkeley, Braxton, Cabell, Fayette, Gilmer,
Grant, Greenbrier, Hardy, Kanawha (5), Marion, Mercer, Mineral, Mingo,
Monongalia, Pendleton (11), Pocahontas, Preston, Putnam, Randolph, Summers,
Wayne, Webster (3), Wetzel, Wood; V-24 to IX-19.

S. aphrodite aphrodite (Fabricius )—Fayette, Greenbrier, Hampshire, Kanawha
(5), Monongalia, Pendleton (11), Preston, Pocahontas, Randolph, Tucker,
Webster (3); VI-3 to X-9.

Euptoieta claudia (Cramer )—Berkeley, Monongalia, Pendleton (11), Preston,
Randolph; VII-5 to IX-26.

Danaidae

Danaus plexippus plexippus (Linnaeus )—Berkeley, Kanawha, Marion, Monon-

galia, Preston, Putnam, Randolph, Webster (3); V-28 to X-10.
Satyridae

Lethe anthedon (Clark )—Berkeley, Boone, Gilmer, Grant, Greenbrier, Hampshire,
Kanawha (5), Mason, McDowell, Mineral (14), Monongalia, Pendleton (11),
Ritchie; V-24 to IX-12.

L. eurydice eurydice (Johansson )—Greenbrier, Kanawha (5); VI.

L. appalachia Chermock—Fayette, Greenbrier (14); VII-7 to VIII-13.

Euptychia gemma gemma (Hubner )—Boone, Kanawha (5, 8); VI-11 to VII-8.

E. hermes sosybius (Fabricius )—Boone; VI to VIII-24.

E. cymela cymela (Cramer )—Berkeley, Braxton, Brooke, Cabell, Fayette, Gilmer,
Grant, Hancock, Hardy, Marion, Mason, McDowell, Mercer, Mineral, Mingo,
Monongalia, Preston, Randolph, Wayne, Wetzel; IV-23 to VIII-13.

Cercyonis pegala pegala (Fabricius )—Barbour, Greenbrier, Hampshire, Hardy,
Kanawha (5), Mercer, Mineral, Monongalia, Nicholas, Pendleton (11), Poca-
hontas, Randolph, Ritchie; VI-25 to IX-3.

ACKNOWLEDGMENTS

The authors would like to express gratitude to all of those persons cited
above who have contributed their knowledge and their collection data,
enabling this list to be the most complete and comprehensive summary of
West Virginia Rhopalocera to date. We are especially grateful to H. K.
Clench for suggestions and review of the manuscript.

INDEX

Burns, 1964

i.
2. Burns, 1976, Personal Communication with Drees
3. Brooks, 1905

VoLUME 32, NUMBER 3 205

4, Clench, 1972

5. Edwards, 1868, 1884, 1897

6. Field, 1941

7. Holland, 1913

8. Irwin, 1972

9. Klots, 1951

10. Lepidopterists’ News, Field Reports, 1951—Preston

11. Lepidopterists’ News, Field Reports, 1953—Nicolay

12. News of the Lepidopterists’ Society, Field Reports, 1974—Drees, Lavy, Parshall

13. News of the Lepidopterists’ Society, Field Reports, 1975—Showalter

14. News of the Lepidopterists’ Society, Field Reports, 1976—Allen, Boscoe, Butler,
Drees, Jensen

15. Observations only

LITERATURE CITED

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Burns, J. M. 1964. Evolution in skipper butterflies of the genus Erynnis. Univ.
Calif. Pub. in Entomol. 37. Univ. Calif. Press, Berkeley and Los Angeles. 216 pp.

Carpe, R. T., A. M. SHapro & H. K. CLENcH. 1970. Sibling species in the eurydice
group of Lethe (Lepidoptera: Satyridae). Psyche 77(1): 70-103.

Criark, A. H. & L. F. Ciarx. 1951. The butterflies of Virginia. Smithsonian Misc.
Coll. 116(7), 227 pp.

CiencH, H. K. 1972. Celastrina ebinina. A new species of Lycaenidae (Lepidop-
tera) from the eastern United States. Ann. Carnegie Museum 44: 33-44.

1975. Boloria toddi or bellona? (Nymphalidae). J. Lepid. Soc. 29(3): 162.

CovELL, C. V., Jk. 1967. A checklist of the butterflies and skippers of Virginia.
J. Lepid. Soc. 18: 21-24.

1974. A preliminary checklist of butterflies of Kentucky. J. Lepid. Soc.

28(3): 253-256.

& G. B. Srratey. 1973. Notes of Virginia butterflies with 2 new state
records. J. Lepid. Soc. 27(2): 144.

Dos Passos, C.F. 1964. A synonymic list of the nearctic Rhopalocera. Mem. Lepid.
Soc. 1. 145 pp.

1969. A revised synonymic list of the nearctic Melitaeinae with taxonomic

notes (Nymphalidae). J. Lepid. Soc. 23(2): 115-125.

1970. A revised synonymic catalogue with taxonomic notes of some nearctic
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Epwarps, W. H. 1868, 1884, 1897. The Butterflies of North America. Vol. 1, 2,
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Fates, J. H. 1974. Checklist of skippers and butterflies of Maryland. Chesapeake
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and Cynthia (Lepidoptera, Nymphalidae). Smithsonian Contributions to Zoology.

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FREEMAN, H. A. 1973. A review of the Amblyscirtes with the description of a new
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Howe, W. H. 1975. The Butterflies of North America. Doubleday. Garden City.
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Irwin, R. R. 1972. Further notes on W. H. Edwards specimens in Illinois museum
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Tue 1974 Fretp SEASON SuMMaARY. 1975. News Lepid. Soc. No. 2-3.

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Woon, C. E. & C. W. GorrscHaLkx. 1942. The Butterflies of Roanoke and Mont-
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Journal of the Lepidopterists’ Society
32(3), 1978, 207—220

PHENOLOGY AND DIVERSITY OF A BUTTERFLY
POPULATION IN SOUTHERN ARIZONA

GEORGE T. AUSTIN

Department of Biological Sciences,
University of Nevada, Las Vegas, Las Vegas, Nevada 89154

ABSTRACT. Butterfly populations were examined on the Santa Rita Experimental
Range in southern Arizona in 1970-71. Thirty-nine species were recorded of which
14 were considered common and typical of the desert grassland community. Butterfly
activity was noted in all months except November 1970 and January and February
1971. Peak species number and abundance were after the summer rains. Reduced
abundance in spring 1971 was attributed to the dryness of the previous winter. Season
lengths for the two years were similar. Univoltinism was 33% of the fauna in 1970
and 46% in 1971 and was a rainy season phenomenon. Species diversity (H’) closely
paralleled species counts.

It is widely recognized that flight periods of butterfly species in most
areas are seasonal depending on several factors including emigration,
voltinism, competition and tolerances of the component species to various
environmental extremes. Differences in interspecific dates of emergence
and lengths of flight periods and intraspecific synchrony result in changes
in diversity throughout the total favorable season. Detailed studies of
phenology and seasonal changes in diversity of butterfly populations in
limited areas are nearly lacking (e.g., Emmel and Emmel 1962, 1963a,
Shapiro 1975). I made observations on relative abundance and phenology
of the butterflies in a small area in southern Arizona from May 1970 to
November 1971, incidental to my studies of bird populations in the same
area. In this paper I examine seasonal patterns of abundance and diversity
of the butterflies and relate these, as possible, to environmental variables.

Study Area

All observations reported here were made on an area of approximately
50 ha. on the Santa Rita Experimental Range, elev. 1150 m, ca. 10 km SE
of Sahuarita, Pima Co., Arizona. The study area, on the western slope of
the Santa Rita Mountains, sloped slightly to the northwest and was dis-
sected by several washes. This resulted in a heterogeneous habitat, typical
of the surrounding area, with the larger trees and shrubs tending to be
concentrated along water courses. The community including the study
area was described as a desert-grassland biotic community (Lowe 1964)
invaded by woody growth due mainly to protection from fire (Humphrey
1968). The dominant vegetation included mesquite (Prosopis juliflora),
paloverde (Cercidium microphyllum), hackberry (Celtis pallida) and

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VOLUME 32, NUMBER 3 209

cholla cacti (Opuntia fulgida and O. spinosior) with an understory of
several grass species, some small woody bushes (especially Acacia
greggii), succulents and herbs.

Rainfall and temperature for the study period are given in Table 1.
Temperatures averaged near normal during the study. Rainfall was near
normal through July 1970; August was drier than average and September
wetter. The winter (October—March) of 1970-71 was one of the driest
on record with rainfall for the period 13.8 cm below average. Thereafter,
1971 rainfall was about average.

Prosopis and Acacia began leafing and blooming in April. Celtis put
out a few leaves at this time and a few grasses and annuals appeared.
The summer rainy season initiated the annual growth of many grasses
and herbs. Celtis and Cercidium came into full leaf at this time and
additional leaf growth and flowering was seen on Prosopis and Acacia.

METHODS

Observations were made by counting all butterflies seen within 5 m
of me as I walked through the area. Most actual counts were made over
a 3-4 hour period between 0700 and 1300 on clear, windless days. Each of
these days was considered as a standard observation day and relative
abundance was expressed as number observed per day by 10 or 11 day
periods. I made observations on other days of species composition which
occasionally added to the species list for that time period. Specimens of
most species observed were obtained from or near the study area.

RESULTS
Species Composition

A total of 39 species (32 in both 1970 and 1971) were recorded during
this study (Table 2). Thirteen species were represented by no more than 2
observations in each of the two years. Nine additional species (D.
plexippus, E. claudia, S. melinus, H. isola, C. eurytheme, N. iole, P.
protodice, P. communis, S. ceos) were seen 2 or less times in one
of the two years. Of the latter, all except H. isola and P. protodice
were recorded in considerably greater numbers in the alternate year.
Three species (M. leda, E. amyntula, C. hippalus) recorded several
times in 1970 were absent in 1971. This leaves 14 species which I consider
as typical and relatively common in this plant association. Nine of these
(A. leilia, D. chara, L. bachmanii, A. palmeri, L. marina, H. ceraunus, E.
nicippee, P. catullus, E. funeralis) are definitely resident in the area;
the others (D. gilippus, V. cardui, C. cesonia, P. sennae, B. philenor) are

aS)
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Oct.

Sept.

Aug.

July

anta Rita Experimental Range, Arizona (X indicates record for species

June

May

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April

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on a day when numerical data were not collected ).°

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TABLE

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212 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

strong flying and wide ranging species which may or may not have used
the area for reproduction.

All species recorded during this study were previously seen by me in
other similar habitat and in other vegetation types in southern Arizona.
No species not recorded on the study area were seen in adjacent areas.
I thus consider the species list relatively complete.

The butterflies of the study area were representative of the Southeastern
Arizona desert scrub habitat with considerable influence from the foothill
canyon habitat as outlined by Brown (1965). All species seen on the
study area were recorded in one or both of the above habitats by Brown
(1965) except V. annabella and E. amyntula.

The Santa Rita Range butterfly fauna belong to 10 families (if
Apaturidae and Heliconiidae are separated from Nymphalidae). Nym-
phalidae and Hesperiidae were represented by the most species (8 each)
followed by Lycaenidae (7) and Pieridae (6). Overall, the familial com-
position was more diverse than in many temperate communities (e.g.,
Emmel and Emmel 1963b) which often lack representatives of families
with southern distribution (Heliconiidae) or with restricted larval food
habits (Apaturidae and Libytheidae).

Apaturidae was by far the most abundant family in terms of individuals,
due to the high abundance and long flight period of A. leilia. Lycaenidae
and Libytheidae were next in individual abundance followed by
Riodinidae and Pieridae. The two families with most species were
represented by very few individuals. This distribution of individuals by
family contrasts sharply with that in the Sierra Nevada of California
where individual and species distribution was similar (Emmel and Emmel

1963b).

Phenology

Activity by adult butterflies was observed in all months except November
1970 and January and February 1971. The general pattern of activity
was similar during the two years (Fig. 1). During spring and early
summer there was a plateau in number of species which then increased
rapidly to a peak in late summer before decreasing to a small number of
species in fall. In both years, there were peaks of abundance in both
spring and late summer with a period of nearly no activity between
(Fig. 1). In 1970, the spring peak was much larger, and the summer peak |
began earlier and lasted longer than in 1971.

Phenology of the fauna as a whole appeared largely related to rainfall.
Precipitation for winter 1969-70 was near average, and the spring popula-
tion in 1970 was probably typical for the area. The very dry winter of

VOLUME 32, NUMBER 3 ANS

00

(0)
oe)

Oo
e)

p
(e)

20

NO. OF INDIVIDUALS

NO. OF SPECIES

4 1970

4- |971

RAINFALL (CM)

M A M J J A S O N D
MONTH

Fig. 1. Rainfall pattern (1 May—5 Sept.) and butterfly activity on the Santa Rita
Experimental Range, Arizona, in 1970 and 1971.

214 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

1970-71 was probably responsible for the reduced diversity and abun-
dance in spring 1971.

Differences in the overall summer flight period can be directly attributed
to differences between the summer rainfall patterns (Fig. 1). The first
heavy rain of 1970 was on 29 June and was followed by another similar
rain less than a week later. These were sufficient to stimulate the summer
growth of vegetation. Butterfly diversity and abundance increased rapidly
after these rains but diversity decreased rapidly during the below average
dryness of August. Abundance increased in September due mainly to the
emergence of a large brood of L. bachmanii. Both diversity and abun-
dance decreased rapidly in late September and early October.

The 1971 rainy season began on 2 July with a relatively light rain. The
next rain did not occur until 14 July after which there were several addi-
tional, closely spaced rains (Fig. 1). The vegetation responded minimally
to the first rain but much of the new growth dried considerably between
rains. The second and subsequent rains initiated and maintained the
usual spectacular summer growth of vegetation. There was no emergence
or immigration of butterflies with the first rain. The second rain, however,
was followed by a rapid increase in species number and abundance similar
to the increase after the first rain of 1970. The timing of the autumn de-
crease was similar to 1970.

The 3 principal spring species showed a peak in about late May or
early June with continued emergence into early fall as noted above.
Several patterns were evident among the rainy season species. Certain
resident species showed an immediate emergence within a few days of
the first rain. These included A. leilia, D. chara, P. catullus and possibly
E. funeralis. Other species including the immigrant D. gilippus, P. sennae
and possibly C. caesonia and the resident E. nicippee occurred in peak
numbers 2-4 weeks after the beginning of the rains. The second brood
of A. palmeri was timed similarly. Additional immigrants, V. cardui and
B. philenor, showed peak abundance about 2 months after the beginning
of the rains. In both years, however, L. bachmanii reached peak abun-
dance in mid-September. In all other instances where sufficient records
exist, the summer peak in abundance was later in 1971 than in 1970
similar to the overall pattern for all species combined as discussed above.
Nearly all species had greater peak abundance in 1970 than in 1971.

The relationships of the various species within a larval food plant guild
are complex. Adults of the principal Prosopis feeders, A. palmeri and
H. ceraunus, were spring fliers with the latter flying slightly later than
the former although A. palmeri had a relatively large brood after the rains
in 1970. Another Prosopis feeder, M. leda, emerged only after the rains

VOLUME 32, NUMBER 3 AUS

began. Of the two species which feed as larvae on Cassia, E. nicippee was
resident and P. sennae was an immigrant. Both showed peak abundance
from early August to mid-September. There was almost complete overlap
of peak flight activity in 1970 but P. sennae showed peak abundance
earlier than E. nicippee in 1971. The Celtis feeding guild, represented by
A. leilia and L. bachmanii, showed overall peak abundance in mid-
September. The two years differed with little overlap in peak flight
period in 1970 and almost complete overlap in 1971.

The few notes taken on adult resource use indicate use of several
flowering species. The spring species were most often seen visiting Acacia
and Prosopis flowers. In summer, flowers of Mirabilis multiflora (used
principally by P. catullus), Ipomoea coccinea (B. philenor), Hymenoclea
salsola (D. gilippus, L. bachmanii) and Zinnia pumila (D. chara, E.
claudia) were visited.

Temporal partitioning may be partly responsible for increased diversity
of some butterfly populations (Clench 1967). Replacement of one set of
species by another within a season allows increased diversity without
increasing competition for adult resources. It must be recognized, how-
ever, that the observed phenology of adult activity is often closely
integrated with the phenology of immature stages which is largely beyond
the scope of the present discussion.

The numerical data in Table 2 were analyzed using the methods of
MacArthur (1964) and Ricklefs (1966). Season length was calculated
using the information-theoretical measure (H’) for the combined abun-
dance of all species (total season) and for each species that was observed
in 4 or more 10 day periods (specific season). The specific seasons were
averaged and divided into the total season resulting in a figure which
indicates species turnover through the total flight season. Use of H’
places greater weight on periods of abundance and less on periods of rarity
and more accurately reflects the length of the flight period than extreme
dates.

Total season length was about 11 ten-day periods in both years (11.4 in
1970, 10.7 in 1971) and average specific season was over 5 ten-day periods
(5.31 in 1970, 5.26 in 1971). There appeared, therefore, to be two turn-
Overs among the species present. The turnover in 1970 was obvious with
the rainy season species clearly replacing the spring species. The 1971
replacement was less clear but apparently involved the replacement of
early rainy season species by those of the later parts of rainy season.
While this may have had some effect in 1970, it was largely marked by the
large spring broods and longer rainy season of that year.

In general, the phenological patterns were similar to those found by

216 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Brown (1965). The major differences were that Brown did not find spring
broods of A. palmeri, L. marina and H. ceraunus, although he found L.
bachmanii and D. chara in spring, two species which I did not observe.
In southwestern New Mexico, Ferris (1976) also found phenology to be
relatively similar to southern Arizona. He noted a spring brood of all
of the above 5 species.

Voltinism

Voltinism varied both inter- and intraspecifically (Table 2). Two (D.
chara, P. catullus) of the 9 common residents were univoltine during both
years. E. nicippe and E. funeralis appeared to be so in 1971 but were at
least bivoltine in 1970. Eight rare species were encountered frequently
enough to determine voltinism. In 1970, M. leda, P. communis and C.
hippalus were univoltine whereas S. melinus, E. amyntula and N. iole
were bivoltine. In 1971, L. eufala was univoltine and H. isola was
bivoltine. Thus 33% of the resident fauna was univoltine in 1970 and 46%
in 1971. Univoltine species, in all cases, appeared in summer after the
beginning of the rains. Univoltinism in other long summer faunas is
principally a spring and early summer phenomenon (Shapiro 1975). The
mid- to late summer univoltinism in southern Arizona is undoubtedly
related to the greatly increased suitability of the habitat following the
summer rains.

The remainder of the Santa Rita fauna was at least bivoltine. Three
species (A. palmeri, L. marina, H. ceraunus ) had large spring and smaller
summer broods, especially in 1971. The summer populations were repre-
sented by scattered individuals observed over a period of 2-3 months.
In 1970, A. palmeri may have had 4 broods. These 3 species may be
spring univoltines in years when the summer rains fail.

Four rare species (S. melinus, E. amyntula and E. funeralis in 1970 and
H. isola in 1971) had a distinct brood in early June and at least one other
of about the same size following the beginning of the rains. S. melinus and
H. isola appeared to have but one post-rain brood; the others possibly 2
or 3. In both years, small numbers of A. leilia occurred before the rains.
There were 3 distinct peaks possibly indicating 3 additional broods after
the rains in 1970, but in 1971 there was but one large peak in mid and
late September. Fresh individuals were noted throughout the flight season
of both years indicating continued emergence.

Ikmergence was limited to after the rains in L. bachmanii, E. nicippe
and N. iole. L. bachmanii had a small brood immediately after the be-
ginning of the rains, a large brood in September and possibly another
one or two smaller broods still later in the season. E. nicippee appeared

VOLUME 32, NUMBER 3 J2AT

2-576 1970

2.0 e (971

0.8

0.4

0:2
A M J J A S O N

MONTH

Fig. 2. Seasonal changes in species diversity (H’) and equitability (J’) of butterflies
on the Santa Rita Experimental Range, Arizona.

to have at least 2 broods in 1970 but probably only one in 1971 unless
the October records were of an additional brood. N. iole had at least 2
broods in 1970 but was all but absent in 1971.

Two immigrant species (D. gilippus, P. philenor) occurred in small
numbers before the rains and reached a definite peak in numbers in
late summer. P. sennae also showed a peak in abundance after the rains.
Other immigrant species occurred irregularly in low numbers after the
rains. Unworn individuals of these species occurred throughout the
season.

218 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Species Diversity

Diversity of a population can be examined in several ways. The method
most often used for butterflies is species counts for each of several seasons.
This diversity measure for the Santa Rita fauna was examined previously
(Fig. 1). Another measure of diversity based on information theory (H’)
accounts for both species numbers and relative abundance. This measure
has not been widely used for insect populations (Janzen and Schoener
1968) mainly because of the wide range of detectability of the component
species (see Shapiro 1975). In this study, I made a concerted effort to
obtain accurate counts in a relatively narrow transect and none of the
species encountered were particularly secretive. I believe that no species
was grossly under- or over-represented in my counts and that the numbers
in Table 2 reflect the true relative abundance. Therefore, I used these
values to calculate H’ for each 10 day period of the two seasons. The ratio
of H’ to maximum diversity possible if each species were equally abun-
dant is another component of diversity termed equitability (J’, see Pielou,
1966). These were also calculated for each 10 day period.

Diversity, measured by H’, showed a seasonal pattern which closely
paralleled species counts (Fig. 2). Linear regression analysis showed
that the number of species explained over 80% of the variation in H’
(r = .899, N = 31). In contrast, equitability fluctuated widely, showed no
correlation (r = .005) with species counts and was most constant during
the rainy season (Table 2). Considering each season as a whole, H’ and
J’ were slightly greater (2.34 and 0.67) in 1970 than in 1971 (2.20 and
0.63). The average J’ for Santa Rita butterflies of about 0.65 is slightly
lower than the average of 0.74 calculated for insect populations containing

a larger number of orders and thus greater trophic diversity (see Austin
and Tomoff, in press).

CONCLUSIONS

The complexity of the phenology of the southern Arizona butterfly
fauna was first indicated by Brown (1965). He noted that most lowland
species were rainy season fliers or had a spring brood and additional
broods during the rainy season. He further recognized the wide annual
fluctuations with populations dependent on the local precipitation.

Several factors seem apparent in the flight patterns of the species
present. Strict seasonal phenology with only the magnitude affected by
rainfall was exhibited by the 3 principal spring species and by L. bach-
manii in fall. Spring flight by the Prosopis feeders is timed for the larvae
to take advantage of the fresh herbage or flower buds and for the adults’

VOLUME 32, NUMBER 3 219

nectar source which appears to be mainly Prosopis and Acacia. The
second brood of A. palmeri appears timed to the new herbage growth by
Prosopis following the summer rains. Flowering by Prosopis following the
rains is limited and this may account for the very small numbers of L.
marina and H. ceraunus at this season.

The flight seasons of the remaining species are nearly limited to after
the beginning of the summer rains. This suggests that rainfall itself
and/or the resultant increase in humidity combined with the warm
summer temperatures act to break diapause of the resident species. The
rapidity of which adults appear after the first rain may relate to the
stage of the life cycle at which diapause occurs or to a cumulative effect
of successive rains. The enforcement of diapause by heat and aridity in
desert regions has been previously noted ( Wiltshire, 1956). Non-resident
individuals of wide ranging species may be attracted by increased vegeta-
tion growth and flowering by many plants. Adults of 12 species of butter-
flies were observed feeding at the flowers of 2 plant species, Mirabilis
and Zinnia, which flower abundantly and nearly exclusively after the
rains begin. The nearly consistent lateness of the 1971 season compared to
1970 (Table 1) is further indication that rainfall is the important ultimate
factor in the phenology of many species.

The seasonality of the Santa Rita Range butterfly fauna differs con-
siderably from other temperate zone faunas previously examined. The
general trend in most populations is for a rather rapid increase in species
number to a mid summer peak and then a decrease into autumn (Shapiro,
1975). Only the tidal marsh in California showed a peak in species
numbers in late summer or fall. No temperate butterfly community
showed the strict dependence on summer rains. These differences in
phenology (and voltinism) can be largely attributed to climatic differ-
ences; the areas studied by Shapiro (1975) were largely Mediterranean
or Mediterranean-montane in climate. The Arizona community was
similar to others, however, in that peak populations corresponded to peak
vegetation growth. Seasonal phenology in southern Arizona shows certain
similarities to that in the Neotropics where Ebert (1969) found peak
activity during the rainy season.

ACKNOWLEDGMENTS

This study was conducted while I was funded to study birds by the
US/IBP Desert Biome Program under National Science Foundation Grant
GB15886 at the University of Arizona. I thank Donald Thomas and
Robert Ricklefs for critical comments on an early draft of the manuscript.

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JOURNAL OF THE LEPIDOPTERISTS SOCIETY

LITERATURE CITED

Austin, G. T., & C. S. Tomorr. Relative abundance in bird populations. Amer.
Natur., in press.

Brown, K. S., Jr. 1965. Some comments on Arizona butterflies (Papilionoidae).
J. Lepid. Soc. 19: 107-115.

Ciencu, H. K. 1967. Temporal dissociation and population regulation in certain
Hesperine butterflies. Ecology 48: 1000-1006.

Epert, H. 1969. On the frequency of butterflies in eastern Brazil, with a list of
the butterfly fauna of Pocos de Caldas, Minas Gerais. J. Lepid. Soc. 23, supple-
ment no. 3.

EMMEL, T. C., & J. F. EmmMen. 1962. Ecological studies of Rhopalocera at Donner
Pass, California. I. Butterfly associations and distributional factors. J. Lepid.
Soc. 16: 23-44.

& 1963a. Ecological studies of Rhopalocera at Donner Pass, Cali-

fornia. II. Meteorological influences of flight activity. J. Lepid. Soc. 17: 7—20.

& 1963b. Composition and relative abundance in a temperate zone
butterfly fauna. J. Res. Lepid. 1: 97-108.

Humpurey, R. R. 1968. The desert grassland. Univ. of Arizona Press, Tucson.

Lowe, C. H. (ed.). The vertebrates of Arizona. Univ. of Arizona Press, Tucson.

MacArruur, R. H. 1964. Environmental factors affecting bird species diversity.
Amer. Natur. 98: 387-397.

PreLou, E. C. 1966. The measurement of diversity in different types of biological
collections. J. Theoret. Biol. 13: 131-144.

RickteFrs, R. E. 1966. The temporal component of diversity among species of birds.
Evolution 20: 235-242.

SHAprro, A. M. 1975. The temporal component of butterfly species diversity. Pp.
181-195 in M. L. Cody and J. M. Diamond (eds.) Ecology and Evolution of
Communities. Belkaap Press, Cambridge, Mass.

Wixtsuire, E. P. 1956. Notes on the diapause of Lepidoptera in hot arid sub-
tropical climates. J. Lepid. Soc. 10: 201-203.

Journal of the Lepidopterists’ Society
32(3), 1978, 221-223

A NEW HINDWING ABERRATION OF CATOCALA
MICRONYMPHA GUENEE FROM KENTUCKY

CHARLES V. COVELL, JR.?
Department of Biology, University of Louisville, Louisville, Kentucky 40208

ABSTRACT. An aberration of the underwing Catocala micronympha Guenée with
forewings like “form gisela” and entirely black hindwings is described from Oldham
County, Kentucky. It is informally named “form sargenti” in honor of Dr. T. D.
Sargent.

Catocala micronympha Guenée was very common in Kentucky in June
and July 1977, and a large number of specimens representing the various
forewing variations was collected in several counties. On the night of
15 June I took at blacklight a single male of what appears to be the first
known individual with forewings of form “gisela” Meyer, but with hind-
wings completely black on the upperside except for the yellowish-white
terminal line, fringe, and apical patch (Fig. 1). The underside of the
hindwing has a diffuse remnant of the median yellow band below the
costal margin (Fig. 3). Wingspan is 4.4 cm. A typical form “gisela” from
Bernheim Forest, Kentucky, also taken on 15 June 1977, is shown for
contrast (Figs. 2, 4).

The locality from which this specimen was taken was the University of
Louisville’s research farm, the Horner Bird and Wildlife Sanctuary, about
20 miles (32 km) southeast of Louisville in Oldham County, near the
hamlet of Brownsboro. No Catocala aberrations of any kind had been
collected or seen there in over 12 years of fairly heavy collecting until this
collection was made. No achromatic hindwing aberrations of C. micro-
nympha were reported in Barnes and McDunnough (1918), Forbes
(1954), or Sargent (1976). Personal communication with Drs. T. D.
Sargent and D. C. Ferguson leads me to conclude that this specimen is

unique.
Sargent (1976) states that, “. . . hindwing polymorphisms are virtually
unknown . . .” in Catocala (p. 77), while the forewings in some species

(such as micronympha) are highly polymorphic. Single examples of hind-
wing aberrations are also rare. He further states (p. 111) that, “In the
Catocala the most prominent “sports” are those involving substantial
alterations of the normally invariable hindwings of a species.” Those
eastern North American underwings for which there are named aberrations
involving all-black or nearly all-black hindwings, where contrasting light-
colored bands are typical, include the following (with descriptions from

1 Univ. of Louisville Contributions in Biology No. 190 (New Series).

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JoURNAL OF THE LEPIDOPTERISTS SOCIETY

Figs. 1-4. Catocala micronympha Guenée. 1. aberration “sargenti” Covell, upper-
side; 2. form “gisela” Meyer, upperside; 3. aberration “sargenti,”’ underside; 4. form
“gisela,” underside.

Sargent): C. muliercula Guenée, ab. “peramens” Hulst (“HW almost
entirely black”); C. ilia (Cramer), ab. “normani’” Bartsch (“blackish FW
from base to pm line, and extended black on HW”); C. unijuga Walker,
ab. “fletcheri” Beutenmuller (“HW entirely black”); C. grynea (Cramer),
ab. “constans” Hulst (“HW almost totally black”); and C. habilis Grote,
ab. “depressans” Sargent, named by him on the basis of a single specimen
with nearly all-black hindwings which escaped his killing jar (Sargent,
1976, p. 111, 113; Plate V, 2).

While Latinized names for aberrations have no standing in zoological
nomenclature, Dr. Sargent coined “depressans” and 3 melanic form names
in his book as convenient “handles” for such forms. I am therefore fol-
lowing his example and name the new aberrant form Catocala micro-
nympha, aberration “sargenti,” in honor of Dr. Theodore D. Sargent in
recognition of his contributions to the study of North American Catocala.

The specimen on which this name is based is now in my possession, but
will be deposited in the U.S. National Museum of Natural History at a
later date,

Note added in proof: On 6 July 1978, Loran D. Gibson collected a second male of

this new form at light at Otter Creek Park, Meade County, Kentucky. The specimen
was in worn condition, and is in the collection of the University of Louisville.

VoLUME 32, NUMBER 3 225

LITERATURE CITED

Barnes, W. J. & J. McDunnoucH. 1918. Illustrations of the North American species
of the genus Catocala. Mem. Amer. Mus. Nat. Hist. 3 (1), 47 p., 22 pls.

Forses, W.T. M. 1954. Lepidoptera of New York and neighboring states. Part III,
Noctuidae. Cornell Univ. Agr. Exp. Sta. Mem. 329, 433 p.

SarcGENT, T. D. 1976. Legion of Night: The Underwing Moths. Univ. of Mass.
Press, Amherst. xii + 222 p., 8 pls.

Journal of the Lepidopterists’ Society
32(3), 1978, 223

CONFIRMATION OF THE OCCURRENCE OF AN ALBINISTIC FEMALE
FORM OF PHOEBIS PHILEA (PIERIDAE) IN EXTREME
SOUTHERN TEXAS

Phoebis philea (Johansson) is a large pierid butterfly common in tropical America.
Individuals from Mexico enter southern Texas often (annually according to Howe,
1975, The Butterflies of North America, Doubleday, Garden City, N.Y., 633 p.). Males
of this species are easily recognized by the striking contrast of yellow and orange
portions of the dorsal forewings and hindwings. Females have the marginal dark
markings typical of females of the genus, with the yellow and orange wing portions
somewhat less contrasting. An albinistic form of philea was named “obsoleta” by
Niepelt (1920, Int. Entomol. Zeit. 14: 17); this form corresponds to albinistic female
forms in other species of Phoebis.

On 12 August 1961 I collected one “obsoleta” in Brownsville, Cameron County,
Texas. The dorsal wing surfaces were quite faded with scales totally lacking in
isolated areas, particularly on the forewing discal cell area. Scales still present tend
to be lightly greenish white. The ventral wing surfaces were also faded, but orange
scales remain in sufficient numbers to provide the general color.

One previous report of “obsoleta” from the extreme southern tip of Texas is known.
Stallings and Turner (1946, Entomol. News 57: 44) reported a specimen collected
in the Lower Rio Grande Valley. H. A. Freeman, who collected this first specimen,
has kindly provided the data as follows: 23 August 1944 at a roadside park between
Pharr and Hidalgo, Hidalgo County. My second specimen is of interest because local
lepidopterists probably are unfamiliar with this form.

Occurrence of philea in southern Texas is seasonal, with most specimens being
reported from September to November (McGuire & Richard, 1974, An Annotated
Checklist of the Butterflies of Bentsen—Rio Grande Valley State Park and Vicinity,
Texas Parks & Wildlife Department, Mission, Texas, 21 p.). The worn condition of
my specimen indicates that it arrived here after long-distance migration from some-
where in northern Mexico. Substantial numbers of philea were found at least as far
north as central Texas in late summer 1971 following an unusual climatic regime
(Neck, unpub. data); no “obsoleta” were seen at this time by local collectors.

RAYMOND W. Neck, Pesquezo Museum of Natural History, 6803 Esther, Austin,
Texas 78752.

294 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Journal of the Lepidopterists’ Society
32(3), 1978, 224

A NEW WEEDY HOST FOR THE BUCKEYE,
PRECIS COENIA (NYMPHALIDAE)

The Buckeye, Precis coenia Hbn., is an opportunistic, oligophagous species hitherto
recorded from several genera of the plant families Verbenaceae, Plantaginaceae, and
Scrophulariaceae in North America. In the Sacramento Valley of lowland central Cali-
fornia it feeds on two species of Lippia (Verbenaceae) and one of Plantago (Plantag-
inaceae) (Shapiro 1974, J. Res. Lepid. 13: 120) while in the nearby Vaca Hills it
occurs on Diplacus (Scrophulariaceae ) and at mid-elevations on the Sierran west slope
on Penstemon azureus Benth. (Scrophulariaceae). In the San Francisco Bay area it
has been found on garden snapdragons (Antirrhinum), an introduced scroph. On 17
September 1977 an infestation of fifth-instar larvae, probably from a single colonization
event, was found on large mats of fluellin, Kickxia spuria (L.) Dumort, growing in
cracks in an abandoned roadway at Davis, Yolo Co., California. This Mediterranean
scroph is closely related to toadflax (Linaria) and to snapdragon and occurs sporad-
ically in lowland California as a pavement and roadside weed. In captivity the larvae
ate leaves, buds, flowers and fruit freely. No recorded hosts of any plant family could
be found within 30 m of the infested plants. Adult P. coenia were present.

Fluellin is a prostrate perennial plant with cordate, dull green, pubescent leaves less
than 1 cm long. The flowers, which are open mostly in the morning, are snapdragon-
like, with a long spur; they are purple and bright yellow and about 1 cm long. The
stems, which may be 50 cm long, form tangled mats up to a meter across.

Artuur M. SHapiro, Department of Zoology, University of California, Davis, Cali-
fornia 95616,

Journal of the Lepidopterists’ Society
32(3), 1978, 224

A SECOND LOCALITY FOR EULYTHIS MELLINATA (GEOMETRIDAE)
IN NORTH AMERICA

While identifying some Nova Scotian moths for James Edsall of Halifax, a specimen
of Eulythis was examined which resembled no species known to occur in the province.
A check on the identity of the specimen at the Nova Scotia Museum showed it repre-
sents a Palearctic species, Eulythis mellinate F. (South, 1972, The Moths of the British ©
Isles, Warne, London, 379 p.), a new provincial record and the second locality in
North America where this moth has been collected. Sheppard (1975, Ann. Entomol.
Soc. Québec 20: 7) recorded this species from Laval, Québec, under the name
associata Borkh,

Eulythis mellinata is a widespread Palearctic species. In Europe and Britain the
larvae feed on red and black currant ( Ribes rubrum L. & R. nigrum L.). Mr. Sheppard
informs me that this species has probably become established on Mountain (Alpine)
currant (Ribes alpinum L.) hedges in the vicinity of his home at Laval, Québec.

The Nova Scotian specimen is a fresh female and was collected on 31 July 1972 at
light in Armdale, Halifax, Nova Scotia. Other captures have been recorded in North

VoLUME 32, NUMBER 3 DNS)

Fig. 1. Eulythis mellinata F. Female from Armdale, Halifax, Nova Scotia. 31 July
1972. J. Edsall. 3.5x.

America at Laval (Isle Jesus), Québec on 10 July 1967 (1 male), 24 June 1973 (1
female), 1 July 1973 (1 male) (Sheppard 1975, Ann. Entomol. Soc. Québec 20: 7),
28 June 1974 (1 male), 7 July 1974 (1 female), 29 June 1975 (1 female), 18 June
1976 (1 male) and 24 June 1976 (1 male) (Sheppard, 1977, pers. comm. ).

The introduction of Eulythis mellinata in Nova Scotia was almost certainly recent
as the specimen was collected in an area which has been intensively collected for the
last 30 years, yet this is the only specimen which has been taken to date. The occur-
rence of the moth in two widely separated localities in eastern Canada indicates well-
established populations, and its occurrence in other eastern North American localities
should therefore be expected. A photograph of the adult has been included to aid in
identification.

KENNETH NEL, Department of Biology, Dalhousie University, Halifax, Nova Scotia.

Journal of the Lepidopterists’ Society
32(3), 1978, 225-226

OCCURRENCE OF THYMELICUS LINEOLA (HESPERIIDAE)
IN NEWFOUNDLAND

The recent rapid spread of the European Skipper, Thymelicus lineola (Ochsen-
heimer) in North America, particularly in the northeastern part of the continent, evi-

226 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

dently has excited considerable interest (Burns 1966, Can. Entomol. 98: 859-866;
Straley 1969, J. Lepid. Soc. 23: 76; Patterson 1971, J. Lepid. Soc. 25: 222). As far as
Canada is concerned it is now listed (Gregory 1975, Lyman Entomol. Mus., McGill
Univ., Ste-Anne de Bellevue, Québec, p. 11) as occurring in the provinces of Québec,
New Brunswick, Nova Scotia, and British Columbia, as well as Ontario, where it was
first noted on this continent in 1910 (Saunders 1916, Ottawa Nat. 30: 116).

The butterfly was certainly present in great numbers in one spot in northeastern
Nova Scotia (Cape Breton Island) on 26 July 1977 where I found it on grassy waste-
land adjacent to an abandoned coal mine at Sydney Mines. This is 3 km north of
Sydney whence the ferry sails for Newfoundland, a voyage of 160 km across the Cabot
Strait. Having arrived in Newfoundland, I found T. lineola in the western part of
the island, on 28 July 1977. The locality was an open grassy area a few metres wide
between woodland and Highway 430, 15 km north of Deer Lake. About a dozen of
the butterflies (all males) were observed, most being fresh specimens. Three speci-
mens were collected and have been deposited in the Can. Nat. Coll., Ottawa.

Holland (1969, J. Lepid. Soc. 23: 33-42) collected in the Deer Lake area in 1965 at
the same time of year and did not report seeing this species; indeed it does not appear
to have been previously reported from Newfoundland. However, the insect has
certainly reached the island now, presumably by traversing the Cabot Strait from Nova
Scotia in the very recent past. It is perhaps possible that this species used the ferry
for the crossing.

W. J. D. Eseruir, P.O. Box 370, 17 Division St., Colborne, Ontario, Canada KOK
IMO.

Journal of the Lepidopterists’ Society
32(3), 1978, 226-228

A PROBABLE NATURAL HYBRID OF PAPILIO EURYMEDON
AND P. RUTULUS (PAPILIONIDAE) FROM IDAHO

Natural interspecific hybrids seem to be as rare among swallowtails as they are
among butterflies in general. In the field the best evidence for hybridization comes
usually from intermediacy of such characters as wing shape and color patterns.

On 18 May 1976, David H. Wagner and I encountered impressive swarms of Papilio
eurymedon Lucas and P. rutulus Lucas visiting muddy spots at the edge of the town
of Lowell, Idaho Co., Idaho. All of the individuals were males. Flying at the same
place but much less common were P. multicaudatus Kirby, P. zelicaon Lucas, Pieris
napi Linné, Anthocharis sara Boisduval, Euphydryas chalcedona Doubleday and
Hewitson, and Celastrina pseudargiolus (Boisduval and LeConte). Some of the
male swallowtail “clumps” on the moist soil included over 50 butterflies. They were
probably seeking sodium (cf. Arms et al. 1974, Science 185: 372-374). Obviously the
situation here was ideal for observing variations, and we examined the crowded butter-
flies carefully in the hope of finding aberrant forms. The differences between P.
eurymedon and P. rutulus were immediately visible as they flew up and settled, often
spreading their wings as they crawled over the moist earth. The gray-white ground
color of the former contrasted with the bright clear yellow of the latter. Also the
much broader black stripes and reduction of ground color of P. eurymedon quickly
separated it from P. rutulus.

In one group of swallowtails we noticed a perplexing individual that did not fit
either P. eurymedon or P. rutulus. Its ground color was whitish lemon-yellow and the

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VoLUME 32, NUMBER 3

Fig. 1. Male swallowtails from Lowell, Idaho. Above, dorsal view; below, ventral
view (magnifications vary slightly). A. Papilio eurymedon. B. Probable P. eurymedon
x rutulus. C. P. rutulus.

stripes were of intermediate width. The odd specimen was captured and is illustrated
together with examples of the two associated species (Fig. 1). The specimen is inter-
mediate in the position, extent, and shape of practically every stripe and spot. This
evidence supports the conclusion that the odd specimen is a natural hybrid between
P. eurymedon and P. rutulus.

228 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

We were especially impressed by the precise intermediacy of the individual.
Generally speaking hybrid butterflies show such intermediacy, but in some cases they
may resemble one parent more than the other. In the genus Limenitis, F: hybrids of
L. archippus Cramer and L. astyanax (Fabricius) are intermediate, but backcrosses
yield both hybrid-like and parent-like morphs (Platt 1975, Evolution 29: 120-141).
In swallowtails at least, even F: hybrids may sometimes show one-sided intermediacy.
For example, Clarke and Sheppard (1957, Lepid. News 11: 201-205) bred female
P. glaucus L., the eastern North American counterpart of P. rutulus, with male P.
eurymedon, and found that the glaucus wing pattern seemed generally dominant to
that of P. eurymedon (cf. their Fig. 2 with Fig. 1 of the present paper). The F:
progeny of laboratory crosses of P. polyxenes and P. xuthus were like the former parent
in 11 out of 14 characters (Remington 1959, J. Lepid. Soc. 13: 151-164). Thus it is
possible that all individuals of P. ewrymedon X rutulus found in nature in the future
will not be so conspicuously and precisely intermediate as the one figured here.

WARREN HERB WAGNER, JR., Department of Botany, University of Michigan, Ann
Arbor, Michigan 48109.

Journal of the Lepidopterists’ Society
32(3), 1978, 228-231

NOTES ON SOME MOSAIC PIERIS (PIERIDAE)

Mosaic specimens occur in many if not all species of butterflies and moths, and are
of scientific interest in that they can provide clues to the sequence of events occurring
in embryonic or post-embryonic development. The origins of several types of mosaics
are discussed by Ford (1945, Butterflies, Collins, London, Ch. 9). The mosaic nature
may be sexual (various kinds of gynandromorphs) or homeotic (involving the produc-
tion of a normal feature or pattern in an inappropriate location) or neither.

The checkered white, Pieris protodice Bdv. & LeC., is one of the most abundant and
widespread North American butterflies and shows a conspicuous sexual dimorphism.
There are apparently no published reports of gynandromorphs or other sexual mosaics
although a bilateral non-sexual aberration inherited in a Mendelian manner has been
reported (Shapiro 1970, Wasmann J. Biol. 28: 245-257). I have never seen a mosaic
in any institutional or private collection. Figure 1 shows the first such specimen I
have turned up in thirteen years of research on this species, including mass laboratory
culture through over 30 generations and repeated field sampling in several states. It
was collected in a sample of 10 taken 16 October 1977 at Rancho Cordova, Sacramento
Co., California. It is a very unusual gynandromorph for a number of reasons. The
entire body and three wings are apparently male. The right forewing appears about
40% female, with the inner margin, discal cell and apex mostly female. There are
three black spots near the margin in the interspaces where no black normally occurs
in either sex. The female characters are confined to the upper surface. Ventrally the
forewings are symmetrical and both male. Thus the assumed chromosomal accident

must have occurred in a cell all of whose progeny were fated to positions on the surface
giving rise to the dorsal lamina and its scales.

I have on hand a similar mosaic gynandromorph of Colias eurytheme Bdvy. (figured
by Shapiro 1973, J. Res. Lepid. 12: 94) in which the apex of the left forewing is
female above, and the rest of the animal male. In this case the sexes do not differ

ventrally in the forewing apical area, and it cannot be said with certainty whether

229

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230 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fig. 2. Mosaic Pieris rapae from California. A: bred, Davis, Yolo Co., April 1972;
B: wild, Davis, March 1972; C: Southport, Yolo Co., March 1973; D: Sacramento
Co., April 1973; E: Suisun Marsh, Solano Co., March 1974.

the specimen is aberrant on both surfaces. The same applies to the Colias philodice
Latr. figured by Emmel (1964, J. Res. Lepid. 3: 63) as a mosaic gynandromorph. In
this case, however, it is unlikely that the specimen is gynandrous at all. The “female”
pattern is chaotic and more likely represents patches of male ground color within the
black border, a not uncommon occurrence.

Figure 2 illustrates some mosaics of Pieris rapae L. from northern and central Cali-
fornia. Two of these are clearly sexual. Specimen A is a bilateral gynandromorph
bred ex ovo at Davis. Specimen B is a rather worn, field-collected specimen. These
are the only California gynandromorphs I have seen in this species, and it is of more

VOLUME 32, NUMBER 3 231

than passing interest that B was collected in the same field in the same week as the
mother of A. Both specimens show some streaky mosaicism, with patches of male
scales on the female side and conversely.

Specimens C through E have abnormal, asymmetrical black markings which are
confined to the upper surface. (Mosaics of this sort are very rare on the ventral surface.
I have only one rapae, a female, with an abnormal black marking confined to the
ventral surface and this is a “ray” similar to and perhaps homologous with the
Mendelian character found in the protodice group (Shapiro 1973, Wasmann J. Biol.
31; 301-311).) Im specimen E the shape of the spot suggests homeosis, but it is
located in an inappropriate interspace; moreover its position corresponds to an obvious
crimp in the dorsal lamina, of the sort to be expected from a deformation of the pupal
wing-case. Such injuries may occur when an unhardened pupa slips in its silken girdle.
In Colias they routinely result in melanization of the area around the injury; if the
adult is able to eclose, the resulting pattern is grossly abnormal (compare female
figured by Shapiro 1970, Entomol. News 81:50/data document p. 5).

Braun (1939, Biol. Bull. 76: 226-240) showed that as pigment precursor spread out-
ward across the wing from the body, deposition took place in those scales which were
sufficiently chitinized at that time. Control of pattern thus depends on the rate of
scale maturation, which may be accelerated around injuries—contributing to mosaics
such as these. Since injuries are most likely on the dorsal surfaces, especially of the
forewings, it is not surprising that mosaicism is commonest there. The black streaking
near the costa of specimen D could have arisen in several ways, but that near the
hindwing apex of C suggests an injury resulting in pigment deposition in the corre-
sponding scales as a wave of melanin precursor moved across the wing.

All of the specimens figured are in the collection of the University of California at
Davis.

ArTHur M., SHaprro, Department of Zoology, University of California, Davis, Cali-
fornia 95616.

Journal of the Lepidopterists’ Society
32(3), 1978, 231-233

A MALE-LETHAL GENETIC FACTOR IN PHYCIODES THAROS
(NYMPHALIDAE)

During the course of four years of rearing studies using Phyciodes tharos Drury, a
total of 17 broods was reared of stock from Upper Tyrone Township, Fayette Co.,
Pennsylvania. Of these, 15 were derived from wild-collected females and the
remaining 2 from wild-laid egg patches found on leaves of the foodplant, Aster simplex.
Exact egg-hatch data were kept on 16 of the broods, viability and sex ratio data on 13.
Of the 17 broods, 3 showed almost total male inviability. In 2 of these 3, most mortality
appeared to be during embryonic development, whereas in the third there was normal
embryonic viability but about 50% mortality between the first and fourth larval
instars. An additional brood (77-63) reared from a wild female collected in Rochester
Mills, Indiana Co., Pennsylvania, showed greatly reduced embryonic viability and an
almost total absence of male adults (Table 1).

Female progeny from two of the abnormal Fayette Co. broods (74-4, 76-2) were
mated to males from normal broods of the same population. Each of these isofemale
lines (A and B) showed a tendency toward lowered egg fertility (P < .001, Wilcox on

932 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TaBLeE 1. Egg fertility, embryonic viability, adult sex ratio, and prepupal and pupal
mortality in broods of Phyciodes tharos from wild-collected eggs and females. “Male-
lethal” broods indicated by asterisks.

Broo No. of Proportion Proportion Total Proportion Proportion
No. Eggs Fertile Hatched Adults Males Mortality
73-1 199 1.000 1.000 U5) 0.493 0.053
73-2 167 1.000 1.000 140 0.464 0.021
13-3* Si 0.978 0.724 207 0.010 0.019
73-4 638 1.000 1.000
73-29 229 1.000 1.000 56 0.428 0.036
73-30 21 0.523 0.381
74-1 162 1.000 1.000
74-2 472 0.998 1.000
74-3 243 0.992 0.984
74-4* 45 1.000 1.000 ~23 0.000 0.000
75-5 161 0.578 0.012
75-7 220 1.000 1.000 Paley 0.524 0.042
75-8 258 0.992 0.976 126 0.484 0.008
75-54 289 0.990 0.996 160 0.519 0.025
75-66 MB) 1.000 0.985 UT 0.416 0.130
15-67 IAS, 1.000 1.000 21 0.762 0.000
76-2* WZ, 1.000 0.444 118 0.000 0.025

77-63* 484 0.983 0.736 174 0.023 0.172

TABLE 2. Egg fertility, embryonic viability, adult sex ratio, and prepupal and pupal
mortality in “male-lethal” isofemale lines of Phyciodes tharos.

Broo No. of Proportion Proportion Total Proportion Proportion
No. Eggs Fertile Hatched Adults Males Mortality

Isofemale Line A

74-15 131 0.191 0.333
74-16 417 0.858 0.640

Isofemale Line B

76-10 106 0.066 0.714
76-11 746 1.000 0.614 126 0.000 0.103
76-13 611 0.988 0.778 9 0.076 0.025
76-14 84 0.060 0.800
76-15 226 0.279 0.714 24 0.083 0.000
76-16 1048 0.995 0.715 150 0.000 0.033

two-sample test) and sharply reduced embryonic viability (P < .001) compared with

the normal broods, Four broods were reared through to adults in Line B. Each of these
showed almost total male inviability (P = .001) (Table 2). In none of the “male-lethal”
broods was there an unusually high incidence of mortality during the prepupal and

pupal stages. The male lethal crisis appears to occur during embryonic or larval devel-
opment.

VOLUME 32, NUMBER 3 230

Robinson (1971, Lepidoptera Genetics, Pergamon, New York, 687 p.) has discussed
male-deficient broods in several species of Lepidoptera. In Abraxas grossularia L.
(Geometridae), a karytotypic aberration in females gives a tendency to produce nearly
unisex but normally viable broods. In Hypolimnas misippus L. (Nymphalidae),
females from some small island populations produce all female broods with reduced
embryonic viability. Here a dominant sex-linked gene has been postulated.

Owen (1966, Heredity 21: 443-451) has investigated East African populations of
Acraea encedon L.. ( Acraeidae ), some of which contained only 0.6 to 6.2% males. Eggs
produced by wild-collected females showed normal viability. Parthenogenesis was
ruled out, and the genetic basis of the unisexual broods remains unknown.

The present case in P. tharos appears to be similar to that in H. misippus. Pre-
sumably, in both cases the disadvantage of heavy selection against male progeny is
offset by some selective advantage to the females carrying the tendency toward uni-
sexual broods.

Cuar.es G. Oxiver, R. D. 1, Box 78, Scottdale, Pennsylvania 15683.

Journal of the Lepidopterists’ Society
32(3), 1978, 233-234

OVIPOSITION BEHAVIOR OF COLONIZED
HYALOPHORA GLOVERI GLOVERI (SATURNIIDAE)

Efficient collection of eggs is an important aspect of maintaining small colonies
of giant silkworm moths as breeding stock. This can be accomplished by establishing
an oviposition profile for the species being reared and collecting eggs only during
the period of peak oviposition. Experience in rearing many species of Nearctic giant
silkworm moths has shown that most eggs are deposited during the first few nights
after mating. Oviposition profiles reported for Hyalophora cecropia (Linnaeus )
(Taschenberg & Roelofs 1970, Ann. Entomol. Soc. Amer. 63; 107-111) and Callosamia
promethea (Drury) (Miller & Cooper, 1977, J. Lepid. Soc. 31: 282-283) are specific
examples of this pattern. This paper reports oviposition data for a small breeding-stock
colony of Hyalophora gloveri gloveri (Strecker) maintained on wild black cherry
(Prunus serotina) in Frederick County, Maryland. Because of the small size of the
colony (<12 individuals) observations were limited to five individuals.

Five female moths, each of which mated on the first night after emergence, were
placed in brown paper bags (lunch size) on the first night after mating; and were
transferred to new paver bags each night thereafter until death. After a period of time
sufficient to allow all eggs to hatch, the bags were opened to record the number of
eggs deposited and the number hatched.

The average longevity of the females after mating was 6.6 days; two individuals
lived for 6 days and three lived for 7 days. The females deposited a total of 776
eggs during the study. The maximum number of eggs deposited by a single female
was 198; the minimum number was 114. The average number of eggs deposited per
female was 155.2. Percent hatch was moderate for eggs deposited during the first 4
nights after mating, the average ranging from 60.7% to 78.4%. The total number of
larvae produced per female ranged from 60 to 137; the average being 103.2. The

234 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

120
100
Z [_] ces peposiTeD
O
=
2 BE Larvae propucen
= ai
Lu
—
=>
Cé
=e
=
© 60
ica)
OS
(Sp)
Liu
bh
S
40
CXS
Lu
=
=
Fz
am
rr
et A)
0

] 2 3 4 5 6 7
NIGHTS AFTER MATING

Fig. 1. Oviposition profile for colonized Hyalophora gloveri gloveri females.

oviposition profile shown in Fig. 1 indicates that H. gloveri gloveri follows the general
pattern reported for other species; and that the optimum time for collecting eggs to
maintain small colonies for breeding stock is during the first night after mating.

THoMas A, Mitier, U.S. Army Medical Bioengineering Research and Development
Laboratory, Fort Detrick, Frederick, Maryland 21701.

(This research was not supported by government funds; the opinions contained herein are those

os the author and should not be construed as official or reflecting the views of the Department of
the Army.)

VoLUME 32, NUMBER 3 | 235

Journal of the pepitentents: Society
S2(3); 1978; 23

THE MURRAY O. GLENN COLLECTION OF MICROLEPIDOPTERA

The collection of microlepidoptera assembled from 1931-1976 by Murray O. Glenn
has contributed significantly to the classification and ecology of the Lepidoptera of
the midwestern prairies in the USA. It includes ca. 30,000 carefully prepared moths
representing 1325 identified species, 949 of which Glenn collected in the prairies,
wooded bluffs, and bottomlands near the Illinois River and its tributaries in Putnam
and Marshall counties in north-central Illinois. Significantly the area is now the type-
locality for 17 species and at least one additional species that currently is being
described.

The species that have been named from the Glenn collection with Putnam County as
the designated type-locality include the following:

OLETHREUTIDAE. Polychrosis sambuci Clarke, Endothenia microptera Clarke,
Exartema comandranum Clarke, Eucosma uta Clarke, Epiblema naomi Clarke, and
Epinotia atristriga Clarke.

COSMOPTERIGIDAE. Teladoma incana Hodges.

MOMPHIDAE. Batrachedra illusor Hodges and Chedra inquisitor Hodges.

WALSHIIDAE. Periploca cata Hodges, Aeaea venifica Hodges, Sorhagenia baucidis
Hodges, and Perimede maniola Hodges.

GELECHIIDAE. Chionodes asema Clarke and Dichomeris glenni Clarke.

OECOPHORIDAE. Agonopterix dimorphaella Clarke.

PTEROPHORIDAE. Oidaematophorus glenni Cashatt.

The holotypes for all the above named species are at the U.S. National Museum of
Natural History (USNM) except for Oidaematophorus glenni which is in the collection
of the Illinois Natural History Survey (INHS).

Glenn, in 1977, donated his private collection to the INHS and the USNM, the
former institution receiving the identified specimens (ca. 20,000) including numerous
paratypes and the latter all unidentified moths (ca. 10,000). (His collection of macro-
lepidoptera was given to the INHS in 1969. )

Glenn, as a collector, “. . . . seemed to have a genius for coming un with rare and
desirable things.” (Klots, pers. comm.). In addition to collecting, he succeeded in
ascertaining natural foodplant associations for many of the species that he encountered.
Much of this information is presently available only by examining the collection itself
but should prove extremely useful to future studies if coupled with the 46-year
compilation of flight records associated with the specimens and environmental changes
induced by natural or artificial factors.

GrorcE L. GopFrey, Section of Faunistic Survey and Insect Identification, Illinois
Natural History Survey, Urbana, Illinois 61801.

_ 1 All specimens collected by Glenn in this defined region are labelled ‘Putnam Co., IL.” However,
it should be noted that while Glenn did most of his collecting in Putnam County he occasionally
forayed into the adjoining portions of Marshall County.

236 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Journal of the Lepidopterists’ Society
32(3), 1978, 236

A NEW RECORD FOR CALYCOPIS CECROPS (LYCAENIDAE)
IN COLORADO BY AIRCRAFT-INTRODUCTION

A new butterfly record for Colorado, Calycopis cecrops (Fabricius), was collected
by Howard Bone, in the rear cargo pit of a United Airlines 727 at Stapleton Inter-
national Airport, Denver, on 12 July 1977. The flight was a charter from the east coast
arriving in Denver around 1030 MDST. The cargo pit doors of such aircraft are left
in the open position the entire time the aircraft is on the ground, and this can easily be
from 30 minutes to over an hour, which is ample time for any insect to fly into the pit.
The mentioning by others (Eliot 1977, J. Lepid. Soc. 31: 75; Riotte 1977, J. Lepid.
Soc. 31: 182) that Lepidoptera can be transported by both military and commercial
airliners is confirmed by this record.

Another possibility of an aircraft-introduced species is that of Tmolus azia
(Hewitson). This species has been collected in the state twice. The first specimen was
collected by Jim Eff on 16 July 1957, Chatauqua Mesa, Boulder Co., Colorado. The
second specimen was collected by Marc Epstein on 26 July 1975, Magnolia Rd.,
Boulder Co., Colorado. These records are both equidistant from the airport. The facts
that 1) there are no other records from the state, that 2) they were collected about the
same time of year, and that 3) there were no other records during the 18 vear interval,
suggests the possibility these two specimens were introduced by aircraft.

This unusual occurrence of stowaways on aircraft could provide a very logical
explanation for the introduction of butterflies at great distances from their normal
ranges. A butterfly could easily fly into the pressurized, air-conditioned cargo pit of an
aircraft, and be flown across the country or overseas in a matter of a few hours.

The normal range of Calycopis cecrops is from eastern Kansas through southern Ohio
to southern New Jersey and southward to Florida and Texas. The closest distance to
Colorado within this range is approximately 500 air miles (805 km), but this specimen
traveled some 1500 air miles (2414 km) from the east coast inside an airplane. The
normal range of Tmolus azia in North America is southern Arizona and southern Texas,
which is approximately 600 air miles (966 km) from Colorado.

The specimens of Calycopis cecrops and Tmolus azia (collected by Marc Epstein )
are deposited in the collection of the Denver Museum of Natural History.

MicHaEL G. PocuEe, Devartment of Zoological Collections, Denver Museum of
Natural History, City Park, Denver, Colorado 80205.

Journal of the Lepidopterists’ Society
32(3), 1978, 236-238

NEW FOODPLANT AND OVIPOSITION RECORDS FOR THE EASTERN
BLACK SWALLOWTAIL, PAPILIO POLYXENES ON AN INTRODUCED
AND A NATIVE UMBELLIFER

Papilio polyxenes (Fabr.) is one of the most common Papilionidae in open fields of
the eastern United States. Its larvae are considered to prefer plants of the Umbellif-
erae, although in laboratory no-choice situations they will eat several species of
Rutaceae, Some individuals can also survive when fed on the cucumber (or mountain
magnolia) tree, Magnolia acuminata L. (Scriber and Feeny, in prep. ).

VOLUME 32, NUMBER 3 ear

In Greene County, Ohio, and Ithaca, New York, the preferred foodplant for
Papilio polyxenes (Fabr.) appears to be the introduced wild carrot, Daucus carota
(L.), as it likely is for most of the northeastern United States. In the eastern United
States, a variety of species of Umbelliferae have been reported as foodplants (Scudder,
1889, The butterflies of eastern United States and Canada, 2: Forbes, 1960, Cornell
University Agr. Expt. Sta. Memoir #371; Teitz, 1972, An index to the described life
histories of Macrolepidoptera of the continental United States and Canada, Vol. 1;
Tyler, 1975, the Swallowtail Butterflies of North America, Naturegraph). In addition
to carrot, those plants upon which polyxenes larvae have been observed naturally in
New York (J.M.S.) are wild parsnip, Pastinaca sativa L., poison hemlock Conium
maculatum L., angelica Angelica atropurpurea L., and goutweed, Aegopodium podo-
graria L. In 1976, polyxenes larvae in Ohio were found (M.D.F.) upon bulb-bearing
water hemlock, Cicuta bulbifera L., angelica, Angelica atropurpurea, and wild parsnip,
Pastinaca sativa in addition to wild and cultivated carrot. Here we have two separate
observations of polyxenes on plants which should be reported due to the apparent
lack of any previous natural observations and also due to their particular ecological
significance.

Our first observation is that of a female ovipositing on a characteristically woodland
native plant species, Cryptotaenia canadensis (L.), Honewort (Fernald, 1950, Gray’s
Manual of Botany, 8th ed.). The observation (J.M.S.) was made on 15 July 1977
at 10:00 hrs along a wooded creek at the end of Carlsbrook Drive in the township of
Beavercreek, Ohio (Greene County). Conditions were favorable for oviposition,
with the temperature approximately 30°C and the humidity also very high. Although
these seem to be the ideal conditions for oviposition of most eastern swallowtails, solar
radiation was probably a very important contributing factor as well, especially with
the potential thermoregulation of body temperature in adults (R. C. Lederhouse,
pers. comm. ).

Although P. polyxenes larvae eat Cryptotaenia in laboratory no-choice conditions
(Erickson, 1975, Psyche 81: 109-130; Scriber, 1975, Comparative nutritional ecology
of herbivorous insects; Generalized and specialized feeding strategies in the Papilion-
idae and Saturniidae Ph.D. Thesis Cornell University, Ithaca, N.Y.), the adults are
rarely seen flying in forested areas where the foodplant occurs. Reasons for use of
this particular wooded patch in Ohio are uncertain. This female may have drifted
into the area more by chance than choice since the wooded habitat was a rather
narrow strip in an otherwise open area of residential lawns and first year successional
habitats. Inside the woods the polyxenes female hovered and circled several Crypto-
taenia plants before depositing one egg in an immature flower head. The female
did not investigate the other Umbelliferae (Sanicula, Heracleum, Osmorhiza) nearby,
and instead flew off across the lawns out of sight. Although larvae ate and survived
upon Heracleum maximum (Bartr.) plants from this same wooded location, post-
flowering Heracleum plants in mid-July were less suitable for larval growth than were
the mid-May plants (Finke, 1977, Factors controlling the seasonal foodplant utilization
by the specialized herbivore, Papilio polyxenes. (Lepidoptera: Papilionidae) M.S.
thesis, Wright State Univ., Dayton, Ohio). We do not know whether the pre-
flowering Heracleum plants would have been more attractive for oviposition by
polyxenes in May. P. polyxenes larvae from Greene County, Ohio refused to eat
Sanicula gregaria Bickni., and others died after several days of eating Osmorhiza
longistylis (Torr.) and O. claytoni (Michx.) (Finke, ibid.).

Utilization of other woodland umbellifer species, Taenidia integerrima and Thaspium
barbinode by Papilio joanae Heitzman may have contributed to reproductive isolation
by habitat and the relatively new species status of this polyxenes relative (Heitzman,
1973, J. Res. Lepid. 12: 1-10). The significance of habitat as an isolating mechanism
for adults and larvae of the two species is an interesting aspect of their ecology which
needs further investigation.

Our second observation was made in Ithaca, New York (Tompkins County) on

938 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

5 June 1977. Eight Papilio polyxenes larvae were observed feeding upon one plant of
the introduced Lovage, Levisticum officinale (Doch). It is uncertain if these larvae
(all of which were molting from the 3rd to 4th instar) were derived from one or
several adults. It is extremely unlikely that they wandered onto the Levisticum from
another plant, as there were no other umbellifers nearby. Larvae were collected
and reared through to pupation on Levisticum. Adults have been preserved as voucher
specimens (at the Univ. of Wisconsin, Madison).

The 1973 and 1974 Ithaca populations of P. polyxenes produced larvae which
consumed Levisticum officinale and grew at rates comparable to those for larvae on
35 other Umbelliferae in laboratory no-choice situations (Scriber, ibid.). Papilio
polyxenes larvae obtained from an adult caught in Costa Rica by Paul Feeny all
refused to eat or else ate and died in a similar no-choice situation (Scriber, pers. obs. ).
Some polyxenes larvae from a Costa Rican female x Ithaca male mating were,
however, able to eat Levisticum and grew to the 2nd and 3rd instar before the culture
was lost to virus (J.M.S. and R. C. Lederhouse). The genetic basis for this feeding
ability remains undetermined. Some recent studies with Costa Rican polyxenes
larvae indicate a marginal and variable ability to utilize Levisticum does exist in this
Central American population (Wm. Blau, pers. comm. ).

The Levisticum officinale—Papilio polyxenes interface would appear to offer a
good system for investigation of the evolutionary dichotomy underlying differences in
ovipositional and larval feeding stimulants. The closely related P. machaon in
Sweden, for instance, will oviposit freely on Levisticum, but 100% of the larvae die
feeding on it (Wilkund, 1975, Oecologia 18: 185-197). Wilkund (ibid) suggests
that the rareness and relatively recent introduction of the plant into Sweden may
partially account for nonavoidance of the plant by ovipositing adults. The differences
in larval feeding success between polyxenes of New York and Costa Rican populations
may also be a function of the amount of time the plant and insect have been in
contact. Levisticum does not, to our knowledge, occur in Costa Rica (Standley, P. C.,
1938, Flora of Costa Rica, Field Mus. Natur. History, Chicago Botanical Series, Vol.
18 (no. 420); and Wm. Blau, pers. comm. ).

In summary, we would like to emphasize the fact that foodplant utilization by
P. polyxenes populations appears variable, depending upon local habitat factors
and plant phenology in any particular year as well as regional or geographic host-
plant preferences which may have evolved over a longer period of time. More field
observations and laboratory studies could clarify many unknown or puzzling aspects
of the coevolution of the Papilionidae and their hostplants.

J. Mark ScriperR AND Mark FINKE, Department of Entomology, University of Wis-
consin, Madison, Wisconsin 53706.

VOLUME 32, NUMBER 3 ZOO

Journal of the Lepidopterists’ Society
32(3), 1978, 239-240

BOOK REVIEW

Tue British BUTTERFLIES, THEIR ORIGIN AND ESTABLISHMENT, by R. L. H. Dennis,
1977. E. W. Classey Ltd., Park Road, Faringdon, Oxon., England SN7 7DR. 318 pp.,
20 figs., 15 tables. $17.50 US.

Here is a book that should be of interest to all students of biogeography. It is
well done but many general collectors of butterflies will find it hard going. First,
it is not a book to help determine what species you have from the British Isles. It
is a book that lives up to its second title. Dennis has divided his book into four sharply
separate sections. North Americans may find the first part, “Geomorphological Frame-
work” a bit puzzling at first. The table on p. 7 (the first page of the first chapter)
sets forth the nomenclature used for the Pleistocene in northwestern Europe, including
the British Isles. Add a third column to this, naming the equivalent North American
terms, and you will be allright.

The other sections in sequence are “Recent Rhopalocera geography and habitat
adjustments,’ “Subspecies and subspeciation,” and lastly “The arrival sequence and
establishment of the British Rhopalocera.” In addition, there are four appendices
containing useful information, particularly for those of us on this side of the ocean
who lack intimate knowledge of the British fauna and flora.

The author has done an unusually good job assembling a wealth of data about the
Pleistocene in the British Isles. He retells this in detail. It is best to have a good
scale map of Great Britain and Ireland at hand unless you are intimately familiar
with the geography of them. Maps would have helped in this section, but I suppose
cost would have been prohibitive. In the second part of the book Dennis treats
two subjects: zoogeography and adaptation to the environment. There is considerable
redundancy, but it is not obtrusive. It may be helpful. Here is clearly demonstrated
why the British Isles is the ideal place for such a study at this time. The region is
essentially a closed system for butterflies with few migrants and substantial sea
barriers. Collectors have been active for two centuries or more and their data are
available. The region is small enough—a total area considerably less than the State of
Montana. The amount of information about the area is greater than that for any com-
parable area in either Canada or the United States. This gives the zoogeographer an
ample working sample. The geography side is equally well-reported. The ideas of
geological mapping and stratigraphy and the foundation of modern geology are
British inventions of the 18th century. Detailed large scale mapping is available for
the entire United Kingdom. All of these are needed before such a task as Dennis
set for himself can be confidently attacked.

The second half of the book, Sections C and D, contains an able discussion of
subspeciation as evident in the British Isles. Dennis’s interpretation of subspeciation,
as related to geography, flora and modern and past climates, can be duplicated no-
where on the American continents. The last section is the interpretation of the data
presented in the first three sections. Here Dennis had two earlier similar studies to
use, and to agree or disagree with. He did all three. B. P. Beirne wrote several times
on the subject and summed up his knowledge in The Origin and History of British
Macrolepidoptera found in the Trans. Roy. Entomol. Soc. Lond., 98, 1947. E. B. Ford
set forth his views in detail in Butterflies published by Collins, London. The latest
edition of this delightful book was released in 1957. Dennis had several advantages
over either of these able authors. He had a large number of precise radiocarbon dates
and the results of the very recent and extensive paleontological studies of Pleistocene
and Recent (Flandrian) insects. Needless to say, Tables 14 and 15, setting forth the
ideas of the three writers, show progressive changes in opinion. It will be a very long

240 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

time before a comparable work can be written for any part of the New World. This
is a beautiful example of what can be done when data are available. The book is
an important guide for those who would engage upon detailed zoogeographic studies.

F. Martin Brown, 6715 S. Marksheffel Rd., Colorado Springs, Colo. 80911.

Journal of the Lepidopterists’ Society
32(3), 1978, 240

OBITUARY
JAMES H. BAKER (1910-1978)

Mr. James “Jim” Huffman Baker, charter member of the Lepidopterists’ Society,
died April 14, 1978 at St. Luke’s Hospital in Boise, Idaho after a long illness. He was
67. Jim was born Aug. 14, 1910 in Baker, Oregon, the son of Deering F. and Bernice
Huffman Baker. He graduated from Baker High School in 1928, was employed by
the Citizen’s National Bank in Baker, and then ran the family grocery, Baker's Super-
market, for over 35 years. Jim was a man of many activities. In addition to his lifelong
interest in insects of many orders, he bowled, traveled, was interested in general nature
study, collected rocks, and was an antique dealer and a gem worker.

He published several scientific papers, and his extensive collecting disclosed several
insects that were subsequently named, including Ewphydryas anicia bakeri Stallings
and Turner, and Celastrina argiolus bakeri (Clench). He worked closely with both
the American Museum of Natural History and the Smithsonian Institution. He was
also a member of the Coleopterists’ Society. |

Jim will be greatly missed by all of his many friends and colleagues who have
enjoyed his company and his family’s hospitality. He is survived by his wife, Ilah;
a son, James Michael Baker; a daughter, Judith Ann Haswell of Pullman, Washington;
and two granddaughters.

J. W. Titpen, 125 Cedar Lane, San Jose, California 95100.

EDITORIAL STAFF OF THE JOURNAL
AusTIN P. Puiattr, Editor

Department of Biological Sciences
University of Maryland Baltimore County, 5401 Wilkens Avenue
Catonsville, Maryland 21228 U.S.A.

Frances S. Coew, Managing Editor
Dovctas C. Fercuson, Associate Editor THEODORE D. SARGENT, Associate Editor

NOTICE TO CONTRIBUTORS

Contributions to the Journal may deal with any aspect of the collection and study
of Lepidoptera. Contributors should prepare manuscripts according to the following
instructions.

Abstract: A brief abstract should precede the text of all articles.

Text: Manuscripts should be submitted in duplicate, and must be typewritten,
entirely double-spaced, employing wide margins, on one side only of white, 8% xX 11
inch paper. Titles should be explicit and descriptive of the article’s content, including
the family name of the subject, but must be kept as short as possible. The first men-
tion of a plant or animal in the text should include the full scientific name, with
authors of zoological names. Insect measurements should be given in metric units;
times should be given in terms of the 24-hour clock (e.g. 0930, not 9:30 AM).
Underline only where italics are intended. References to footnotes should be num-
bered consecutively, and the footnotes typed on a separate sheet.

Literature Cited: References in the text of articles should be given as, Sheppard
(1959) or (Sheppard, 1959, 196la, 1961b) and all must be listed alphabetically
under the heading LrreRATURE CrreED, in the following format:

SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd. ed. Hutchinson,
London. 209 p.

196la. Some contributions to population genetics resulting from the
study of the Lepidoptera. Adv. Genet. 10: 165-216.

In the case of general notes, references should be given in the text as, Sheppard
(1961, Adv. Genet. 10: 165-216) or (Sheppard 1961, Sym. Roy. Entomol. Soc.
London 1: 23-30).

Illustrations: All photographs and drawings should be mounted on stiff, white
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headed EXPLANATION OF FicuRES, with a separate paragraph devoted to each page of
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Tables: Tables should be numbered consecutively in Arabic numerals. Headings
for tables should not be capitalized. Tabular material should be kept to a minimum
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Correspondence: Address all matters relating to the Journal to the editor. Short
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ALLEN PRESS, INC. et LAWRENCE, KANSAS
UusS.m

CONTENTS

Tue INFLUENCE OF ENVIRONMENTAL FACTORS ON ROOSTING IN THE

BLACK SWALLOWTAIL, PAPILIO POLYXENES ASTERIUS STOLL

(PaPILIONDAE). John Edward Rawlins & Robert C. Leder-
house) 2
NoTeEs ON THE LiFE CYCLE AND NATURAL HIsTOoRY OF BUTTERFLIES
or Ex SALvaporE. IIC. SMYRNA BLOMFILDIA AND S. KARWINSKII
(NYMPHALIDAE: CoLosurini). Albert Muyshondt, Jr. & Alberto
Muyshondt 0 ee
SCELIODES LAISALIS (PYRALIDAE): DESCRIPTION OF THE MATURE
LARVA AND NOTE ON ITs FEEDING Hasit. E. O. Ogunwolu ___
MIGRATION AND RE-MIGRATION OF BUTTERFLIES THROUGH NORTH
PENINSULAR FLORIDA: QUANTIFICATION WITH MALAISE TRAPS.
Thomas J. Walker
Hysrios BETWEEN CALLOSAMIA AND SAMIA (SATURNUDAE). Richard

S. Peigler EE
RHOPALOCERA OF WEST VircINIA. Bastiaan M. Drees & Linda
Butler 2 a
PHENOLOGY AND DIVERSITY OF A BUTTERFLY POPULATION IN SOUTHERN
Arizona. George T. Austin a)
A New Hinpwinc ABERRATION OF CATOCALA MICRONYMPHA GUENEE
FROM Kentucky. Charles V. Covell, Jr...

GENERAL NOTES

Confirmation of the occurrence of an albinistic female form of Phoebis
philea (Pieridae) in extreme southern Texas. Raymond W. Neck _..
A new weedy host for the Buckeye, Precis coenia (Nymphalidae). Arthur
M. Shapiro 2
A second locality for Eulythis mellinata (Geometridae) in North America.
Kenneth Neil 2
Occurrence of Thymelicus lineola (Hesperiidae) in Newfoundland. W. J.
D. Eberlie

A probable natural hybrid of Papilio eurymedon and P. rutulus (Papilion-—

idae) from Idaho. Warren Herb Wagner, J. -.-2------2----22----2-eeneeeneeee
Notes on some mosaic Pieris (Pieridae). Arthur M. Shapiro ....-...-----.-
A epee genetic factor in Phyciodes tharos (Nymphalidae). Charles
G. Oltwer on I
Oviposition behavior of colonized Hyalophora gloveri gloveri ( Saturniidae).
Thomas A. Miller 2.0500 OE)
The Murray O. Glenn Collection of Microlepidoptera. George L. Godfrey
A new record for Calycopis cecrops (Lycaenidae) in Colorado by aircraft-
introduction. Michael G. Pogue —....)\
New foodplant and oviposition records for the eastem Black Swallowtail,
Papilio polyxenes on an introduced and a native Umbellifer. J. Mark
Scriber & Mark Fink

Nores AND News
BOOK “REVTew nn DR ALA
Osrrganen ok

145

160

175

178

19]

198

207

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Volume 32 1978 Number 4

JOURNAL

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LEPIDOPTERISTS’ SOCIETY

Published quarterly by THE LEPIDOPTERISTS’ SOCIETY
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28 February 1979

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Cover illustration: Dasychira dorsipennata larva, dorsal and lateral views. From
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\

JOURNAL OF

Tae LeEpiIpopreERIStTs’ SOCIETY

Volume 32 1978 Number 4

Journal of the Lepidopterists’ Society
32(4), 1978, 241-250

THE ZALE SETIPES SPECIES COMPLEX (LEPIDOPTERA:
NOCTUIDAE)

EE Topp

Systematic Entomology Laboratory, IIBIII, Federal Research, Sci. & Educ. Admin.,
U.S. Dept. of Agriculture’

ABSTRACT. The setipes complex of the noctuid genus Zale Hiibner is revised.
The identity of two species, confused for more than 100 years, is clarified. Letis
incipiens Walker is removed from the synonymy of Zale setipes (Guenée) and ele-
vated to a subspecies of Z. peruncta (Guenée). Z. discisigna discisignata Draudt is
cited as new synonym of Z. setipes (Guenée) and Z. setipes 2 f£. postmedialis Draudt,
Homoptera aemona Druce (in part), Zale notipennis Draudt are new synonyms of
Z. peruncta (Guenée). The first United States record of typical Z. peruncta {Guenée )
is listed.

The noctuid genus Zale Hubner, as currently recognized, is composed
of a large number of moderately large moths, many with a rather
similar pattern of cryptic wing maculation. The pattern of maculation
is usually composed of numerous irregular transverse or oblique lines,
the moths presumably resembling the bark of trees on which the moths
may rest. They vary in color from nearly black to pale yellow brown
or light gray. A few species, especially some from tropical America,
have areas of pale green scaling on the wings, but that color usually
fades very rapidly after death to yellow or yellow brown. Identification
of species has been difficult in the past and many misidentifications
have occurred. Two closely related species, Zale setipes (Guenée) and
Z. peruncta (Guenée) have been confused, misidentified and misnamed
since 1869. The purpose of this paper is to indicate the proper appli-
cation of the names, to describe and illustrate the characters that dis-
tinguish the species, to detail the specific geographic distributions and
to record Z. peruncta (Guenée) from the United States (Texas).

1¢/o U.S. National Museum, Washington, D.C. 20560.

242 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

The two species are very similar in maculation and both species are
sexually dimorphic, both males and females of each species more closely
resembling the same sex of the other species than the opposite sex of
their own species. The dimorphism is expressed in differences in the
shape of the forewing and in the pattern of maculation. The males
have a narrower forewing, somewhat produced apically, the termen
nearly straight or even excavate before tornus, therefore differing from
the females and the other species of Zale which have broad forewings
and a rounded termen. The pattern of maculation of the males ap-
proaches that of some species of Metria Hiibner (= Safia Guenée). The
females resemble females of other Zale species in wing shape and
maculation. Examples of setipes are consistently larger and have more
pale scaling in the postmedial area of the forewing than do examples
of peruncta. Excellent characters for specific separation exist in the
male genitalia and in the shape of the sternal plate of the eighth ab-
dominal segment of the female.

In 1965 the author studied the types and syntypes of Metria and Zale
in the collection of the British Museum (Natural History) in order to
correctly identify species from the Antilles and to obtain information
necessary for possible future generic revisions. Nearly 80 slides of geni-
talia, mainly of types, were prepared and the errors in the application
of the names, Zale setipes (Guenée) and Zale discisigna (Walker), were
discovered. The types of all the names relating to the setipes complex,
including the lectotype of Z. peruncta (Guenée) which was sent to me
at that time from Paris, were studied. |

History

Guenée described Xylis setipes (1852, p. 7, Noctuélites Pl. 15, Fig. 6)
from a single male from Nova Friburgo, Brazil and Homoptera peruncta
(1852, p. 9) from 2 specimens without locality. Guenée suggested that
one specimen of peruncta was a male lacking antennae, but this seems
unlikely since he placed the male of setipes in a separate genus, Xylis
Guenée, while placing peruncta in Homoptera Boisduval with other
typical Zale species. The lectotype of peruncta is a female specimen
from the Paris Museum selected by Viette (1951, p. 161). The colored
illustration of the type of setipes accompanying the original description
is excellent.

For a number of years the relationship of setipes and peruncta and
the sexual dimorphism in the complex were not recognized. During
that period Walker described males of peruncta as Homoptera ustipennis
({1858| 1857, p. 1071) and Letis incipiens (1858, p. 1266). In 1869 (p.

VoLuME 32, NUMBER 4 243

157) Herrich-Schaffer identified specimens of peruncta from Cuba (true
setipes is not known from the Antilles) as Xylis setipes Guenée and
the trivial name has since been misapplied by all authors to date. In
the collections of the U.S. National Museum and the British Museum
(Natural History) the name was likewise misapplied. Moschler (1890,
p. 202) listed females in his treatment of “setipes,” but no discussion
of sexual dimorphism was included. He did, however, wonder why
the females he studied were only 4245 mm in expanse whereas the
size given for setipes in the original description was 55 mm. Butler
(1879, p. 41) recognized that Walker’s ustipennis was related to setipes
in the statement: “H. ustipennis, a Xylis.” There is no indication as to
his specific concept of setipes. The sexual dimorphism of the complex
had still not been recognized by Druce (1889, p. 341). He utilized
setipes in the same sense as Herrich-Schiaffer and listed ustipennis as a
separate species from Panama; both names were placed in Xylis. He
did not refer to Homoptera peruncta Guenée. Females of both species
of the complex obviously were present in his series of the new species,
Homoptera aemona, because he stated: “The specimens from Guatemala
are rather larger and are paler in colour than those from the Volcan de
Chiriqui. Our figure is taken from one of these latter.” In an unex-
plained action Hampson (1898, p. 250) placed setipes in Polydesma
Boisduval and used ustipennis as a form of that combination for ex-
amples of peruncta from St. Lucia and Grenada. Hampson (1913, pp.
208-210, text figs. 54 and 55) treated both species and provided keys
to and illustrations of the males. He placed the generic names Homop-
tera Guenée and Xylis Guenée in the synonymy of Zale Hiibner. The
two species of the setipes complex (as subgenus Xylis) were separated
from the other species of Zale in the key because the hind tibiae of
male are fringed with long hair and the hindwing with termen some-
what excurved at middle, the costa lobed [expanded] at the base. The
type of setipes now in the British Museum (Natural History) was not
available to Hampson as it was not received by that institution until
1928. Unfortunately, Hampson apparently did not check the original
description and illustration of setipes and continued to use the name
incorrectly for peruncta which he placed along with all other names of
the complex in the synonymy of setipes as identified by him. Hampson
made another error in treating true setipes by calling it Zale discisigna
(Walker). He thought the worn, damaged female holotype of Ho-
moptera discisigna Walker ( [1858] 1857, p. 1066) represented the female
sex of the large species of the complex. This error was perpetuated in
collections and in the literature. Homoptera discisigna Walker does

244 JoURNAL OF THE LEPIDOPTERISTS SOCIETY

resemble setipes somewhat and cons:dering the condition of the type,
the error is partially understandable, but discisigna is considerably
smaller and is not even congeneric. The type of discisigna had been
studied before by other workers and the species placed in Peteroma
Schaus or Barcita Moschler in collections. Dognin apparently was con-
fused by the use of Homoptera discisigna Walker for different species
in collections and must have written to William Schaus about the
problem. He attached a note from Schaus’ reply of June 22, 1922 on a
male of peruncta from Tucuman, Argentina now in the U.S. National
Museum. Schaus informed him that he thought Walker had described
two species named discisigna and that he believed that the one refer-
rable to Peteroma was described in “Characters of Lep. Het.” Schaus
obviously was wrong; no other description by Walker with the trivial
name discisigna has been located. It seems likely that Schaus’s belief
in a second description probably developed because discisigna was
originally described as a Homoptera and because Hampson applied the
name to a Zale species.

SYSTEMATICS

Zale setipes (Guenee )
(Figures 1-2, 9 and 12)

Xylis setipes Guenée, 1852, p. 7; 1858, Pl. 15 ( Noctuélites), Fig. 6—Walker, 1857,
p. 1052.—Druce, 1889, p. 341 (in part ).—M6schler, 1890, p. 202 (in part).

Zale setipes (Guenée), Draudt, 1940, Pl. 70, row b (setipes @ ).

Xylis ustipennis, Druce not Walker, 1889, p. 342 (in part).—Hampson, 1913, p. 208
(synonym of discisigna, Hmpsn. ).

Homoptera aemona Druce, 1889, p. 344 (in part )—Hampson, 1913, p. 208 (synonym
of discisigna, Hmpsn. ).

Homoptera discisigna, Druce not Walker, 1890, p. 345 (in part ).—Hampson, 1913,
p. 208.

Zale discisigna, Hampson not Walker, 1913, p. 208, Fig. 54——Haimbach, 1928, p.
216.—Draudt, 1940, p. 454 (in part).

Zale discisigna ab. discisignata Strand, 1917, p. 43 (= discisigna ab. 1 of Hampson.
An infrasubspecific name, excluded. ).

Zale discisigna discisignata Draudt, 1940, p. 455 (= discisigna ab. 1 of Hampson and
ab. discisignata Strand.) [New synonymy. ]

Diagnosis. Length of forewing, male, 24 to 27 mm, average 24.8 mm; female,
23 to 27 mm, average 24.2 mm. Pattern of maculation as illustrated (Figs. 1 and 2).
Ground color of male paler than female and males of Z. peruncta (Guenée); trans-
verse lines in medial area of forewing distinctly marked; hindwing of male with dark
subterminal shade between veins M, and Cw, reaching termen only at vein M3.
Maculation of hindwing of female variable, with (Fig. 2) or without blue-white
postmedial spots, ground color sometimes paler than females of peruncta, but usually
about the same darkness. Male genitalia as illustrated (Fig. 9), apical process of
ventral margin of valve rather sigmoid in shape, longer than the thin, rather rec-
tangular apical process of costa of valve. Base of uncus with triangular (apex slightly

VoLuME 32, NuMBER 4 245

6

Adults of Zale setipes complex. Fig. 1, setipes, ¢, Chiriqui, Panama; 2, setipes, 2,
“Cent. Amer.”; 3, peruncta peruncta, ¢, Juan Vinas, Costa Rica; 4, p. peruncta, ©,
Orizaba, Mexico: 5, p. incipiens, 6, Cuba; 6, p. incipiens, 2, Convento, Dominican
Republic.

curved distad) lateral flanges: flanges present also on tegumen, bilobed, the depres-
sion between lobes variable in depth and caudal lobe sharp pointed (Fig. 9) or
shorter and rounded (Noctuidae genitalia slide No. 5033 of holotype). Female
genitalia (Fig. 12) with a pair of large rectangular sternal plates present below
ostium.

Types. The Houoryre, ¢, of Xylis setipes Guenée from Nova Friburgo, Brazil,
Noctuidae genitalia slide No. 5033 and the Hotoryre of Zale discisigna discisignata
Draudt, a 2 from Volcan de Atitlan, [Guatemala], Noctuidae genitalia slide No. 5087,
are in the British Museum (Natural History), London, England.

Distribution. The species is known to occur from Mexico to Brazil, but is not
known from the West Indies. Specimens from the following localities have been
examined. MEXICO: Jalapa; Orizaba. GUATEMALA: Volcan de Atitlan; Chejel:
Cayuga: “Guatemala.” COSTA RICA: Tuis; Juan Vinas. PANAMA: Chiriqui.
COLOMBIA: Pacho, Ost-Cordill. ECUADOR: Jatunyacu, Oriente; Abitagua, Ori-

246 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

ente. BRAZIL: Castro, Parana; Rio Janeiro; Petropolis; “Casa Br.’; Ponte Nova, Rio
Xingu, Amazonas; Nova Friburgo.

Discussion. The larger size, paler ground color and distinctive male and female
genitalia distinguish this species from Z. peruncta (Guenée).

Zale peruncta peruncta (Guenée)
(Figures 3-4, 8 and 11)

Homoptera peruncta Guenée, 1852, p. 9.—Walker, 1857, p. 1069.—Viette, 1951, p.
161 (Lectotype designation. ).

Xylis setipes, Gundlach not Guenée, 1881, p. 358 (in part).—Druce not Guenée,
1889, p. 341 (in part )—Mo6schler not Guenée, 1890, p. 202 (in part).

Polydesma setipes, Hampson not Guenée, 1898, p. 250.

Zale setipes, Hampson not Guenée, 1913, p. 209, fig. 55 (peruncta (Guen.), ustipennis
(Wlk.), incipiens (Wlk.) and aemona (Druce) as synonyms ).—Wolcott not
Guenée, 1923, p. 169; 1936, p. 432; 1951, p. 603.—Haimbach not Guenée, 1928,
p. 216.—Draudt not Guenée, 1940, p. 455 (in part ).—Schaus not Guenée, 1940,
p. 229 (in part ).—Biezanko, Ruffinelli and Carbonell not Guenée, 1957, p. 50.

Zale setipes ab. postmedialis Strand, 1917, p. 43 (= setipes, ab. 1 of Hampson. An
infrasubspecific name, excluded. ).

Zale setipes 2 f. postmedialis Draudt, 1940, p. 455, pl. 70, row b (= setipes, ab. 1 of
Hampson and ab. postmedialis Strand). [New synonymy. ]

Homoptera ustipennis Walker, 1857, p. 1071. [New synonymy. ]

Xylis ustipennis (Walker), Butler, 1879, p. 41.—Druce, 1889, p. 342 (in part).

Polydesma setipes £. ustipennis (Walker), Hampson, 1898, p. 250.

Homoptera aemona Druce, 1889, p. 344, pl. 31, fig. 3 (in part). [New synonymy. ]

Zale discisigna, Draudt not Walker, 1940, pl. 70, row 6 ( é, discisigna).

Zale notipennis (sic) Draudt, 1940, p. 455 (misspelling of ustipennis Wlk. ? As
synonym of setipes, Draudt). [New synonymy. ]

Diagnosis. Length of forewing, male, 19 to 23 mm, average 20.4 mm; female,
19 to 23 mm, average 21.7 mm. Pattern of maculation of male (Fig. 3) similar to
that of setipes, but ground color, particularly median part of forewing darker; dark
subterminal shade of hindwing between veins M and Cu reaching termen for most
shades width. Female marked and colored as in setipes, sometimes slightly darker,
hindwing maculation variable as in setipes. Male genitalia with process from costa
of valve longer than process from ventral margin, the latter slightly clavate or mitten-
shaped, both processes (Fig. 8) quite different than in setipes. Flanges at base of
uncus in typical subspecies nearly rectangular. Flanges of tegumen thornlike, apices
slightly recurved. Sternal plate of female genitalia ovoid, caudal margin variable,
usually terminating in two short bluntly pointed processes with a prominent narrow
medial emargination (Fig. 11), occasionally emargination reduced in length, an ex-
treme example with median caudal lobe that is very weakly emarginate (lectotype of
Homoptera aemona Druce).

Types. The Lecroryre, 2, of Homoptera peruncta Guenée, locality unknown, is
in the Muséum National, Paris, France. The Hotoryre of Homoptera ustipennis
Walker, 4, locality unknown, Noctuidae genitalia slide No. 5091, and the syNTyPEs
of Homoptera aemona Druce and Zale setipes £. postmedialis Draudt are in the British
Museum (Natural History), London, England. Druce had examples of both this
species and true setipes in his original series of aemona from Mexico, Guatemala, and
Panama but did not indicate the number of examples either in total or from the
respective countries. He illustrated a specimen, a 9, from Volcan de Chiriqui, Pan-
ama. There are three specimens in the British Museum (Natural History) from that
locality. One labeled Homoptera aemona Druce, Type @, Noctuidae genitalia slide
No. 5090 has been selected and is presently designated as LECTOTYPE. The name

VoLuME 32, NuMBER 4 QAT

woe

Male and female genitalia of Zale setipes complex. Fig. 7, peruncta incipiens, 4;
8, p. peruncta, 6 aedeagus not shown; 9, setipes, ¢ aedeagus not shown; 10, p.
incipiens, 2; 11, p. peruncta, 2; 12, setipes, 2.

postmedialis was proposed for “ab. 1” of Hampson who did not indicate number of
specimens or locality. The specimen labeled as type, a female, Noctuidae genitalia
slide No. 5088 from Grenada has been selected and is now designated LECTOTYPE.

Distribution. The typical subspecies occurs from southern Texas to Argentina on
the continent and in the Antilles from Grenada to Puerto Rico. The specimen from
Texas was collected on 27 November 1973 by A. and M. E. Blanchard. It represents
a new record for the United States. I have examined specimens from the following
localities. TEXAS: Santa Ana Refuge, Hidalgo Co. MEXICO: Jalapa; Misantla;
Orizaba; Cordoba; San Cristobal las Casas, Chiapas. COSTA RICA: Tuis; Juan
Vinas. PANAMA: Chiriquii COLOMBIA: Sta. Marta. VENEZUELA: Aroa.
ECUADOR: Abitagua, Oriente. BRAZIL: St. Catherines [Santa Catarina]; Alta da
Serra, Sao Paulo; Rio Janeiro; Theresopolis.s PARAGUAY: Sapucay; “Paraguay.”
ARGENTINA: Tucuman. GRENADA: Grand Etang; “Grenada.” ST. VINCENT:
Montreal District. ST. LUCIA: 1.5 mi S. Mt. Gimie; “St. Lucia.” DOMINICA:
Clarke Hall; Grand Savanne; Pont Casse. VIRGIN ISLANDS: Gallows Point, St.
John. PUERTO RICO: 4 mi SE. Ciales; Ciales.

248 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Discussion. This species is smaller than setipes and the ground color is slightly
darker, especially the median area of the forewing and the subterminal spot between
veins Ms and Cus. The two apical processes of the valve of the male gentitalia and
the sternal plate of the female genitalia are differently shaped than those structures
in setipes. This subspecies may be separated from the other subspecies by area of
occurrence and by characters of the male and female genitalia discussed in the
diagnosis of the atypical subspecies. The species of the setipes complex of Zale do
not appear to be common in most areas judging from the number of examples in
collections and from my personal collecting experience. Only at Grand Etang,
Grenada, to my knowledge, has a species of the complex been collected in large
numbers (personal light trap collecting). Time of year, weather conditions, time of
night, collecting locality, and collecting technique may in part explain the reduced
captures elsewhere, but I believe some other factor is responsible. A few days later
on St. Vincent and then on St. Lucia collecting with traps in apparently similar
ecological locations resulted in only a few captured specimens. In a three year survey
of Dominica, 1964-1966, eight different collectors collected only four examples of
peruncta. It is true that traps were not utilized there and the species are also known
to be only temporarily attracted to light, settling soon on the plants some distance
away. However, I personally collected other species of Zale there in large numbers
by collecting on such plants, the specimens logated by their glowing, light reflecting
eyes.

Zale peruncta incipiens (Walker), new status

(Figures 5-6, 7 and 10)

Letis incipiens Walker, 1858, p. 1266.—Hampson, 1913, p. 209 (synonym of setipes,
Hampson ).—Schaus, 1940, p. 229 (synonym of setipes, Schaus ).

Zale incipiens (Walker), Draudt, 1940, p. 455 (synonym of setipes, Draudt).

Xylis setipes, Herrich-Schaeffer not Guenée, 1869, p. 157.—Gundlach not Guenée,
1881, p. 358 (in part); 1891, p. 195.—Druce not Guenée, 1889, p. 341 (in
part )—Moschler not Guenée, 1890, p. 351.—Ragués not Guenée, 1914, p. 141.

Xylis sctipes (sic), Anonymous not Guenée, 1895, p. 73 (misspelling of setipes).

Zale setipes, Hampson not Guenée, 1913, p. 209 (in part)—Schaus not Guenée,
1940, p. 229 (in part).

Diagnosis. Length of forewing, male, 18.5 to 20.5 mm, average 19.5 mm;
female, 20.0 to 22.0 mm, average 21.2 mm. It seems likely that the range in size
will probably approach that of the typical subspecies when more material is available
for study. Only five pairs have been examined. The pattern of maculation appears
to be essentially identical to that of peruncta peruncta and similarly variable. The
male and female genitalia differ consistently from those of the typical subspecies.
The apical processes of the valve of the male genitalia (Fig. 7) are more slender
than in typical peruncta, the process of the costa distinctly sinuous. Flanges at base
of uncus thornlike, each with apex bent caudad. Flanges of tegumen much larger,
not thornlike in shape as in typical peruncta, apex variable in shape, up-curved and
blunt (Fig. 7) or sharp-pointed and caudally directed (holotype). Female genitalia
with sternal plate of eighth abdominal segment smaller than in typical subspecies,
the caudal lobes larger in proportion to plate size (Fig. 10).

Type. The Hororypr, 2, from St. Domingo, Noctuidae genitalia slide No. 5089
is in the British Museum (Natural History), London, England.

Distribution. Known only from Cuba and Dominican Republic. The specimens
studied are labeled as follows. CUBA: Santiago; Cayamas; “Cuba.” DOMINICAN
REPUBLIC: St. Domingo; San Francisco Mts., St. Domingo; Hotel Montana, 10 km
NE Jarabacoa, La Vega Proy.; 1.3 km S Loma de Cabrera, Dajabon Prov.; Convento,
12 km S Constanza.

VoLuME 32, NuMBER 4 249

Discussion. The true status of this entity is not known. It has been placed as a
subspecies of peruncta because of the geographic isolation and to express the close
relationship of the two entities compared to setipes. At the present time incipiens
and typical peruncta occur on the neighboring islands of Hispaniola and Puerto Rico
respectively. The former population probably representing an old invasion from
Central America, the latter a more recent invasion from northern South America
through the Lesser Antilles.

LITERATURE CITED

AnonyMous. 1895. Catalogo numérico del museo zool6gico Cubano (Museo Gund-
lach). Alvarez, Habana. 112 pp.

BrEZANKO, C. M. DE, A. RUFFINELLI, & C. S. CARBONELL. 1957. Lepidoptera del
Unieuay. Revta. fac. Agron. Mioncaidee! No. 46; 1-152.

Butter, A. G. 1879. On the Lepidoptera fof the Amazon, collected bye Dr. James
W. H. Trail, during the years 1873 to 1875. Trans. Sauomall Soc. London, 1879,
pp. 19-76.

Draupt, M. 1939-1940. Noctuidae: Catocalinae. In Seitz, Die Gross-schmetter-
linge der Erde, 7: 417-461, pls. 57-71. Stuttgart: Kernen. [pp. 417-428, 1939;
pp. 429-461, 1940.]

Druce, H. 1881-1900. Lepidoptera: Heterocera, Vol. 1. In Godman & Salvin,
Biologia Centrali-Americana, Zoologia, Insecta. i—xxxii, 1-490, pls. 1-41. Lon-
don: Taylor & Francis. [pp. i-xxxii, 1900; pp. 1-24, 1881; pp. 25-32, 1883; pp.
33-112, 1884: pp. 113-160, 1885; pp. 161-200, 1886; pp. 201-256, 1887; pp.
257-344, 1889; pp. 345-440, 1890; pp. 441-490, 1891.]

GuENEE, A. 1852. Histoire Naturelle des Insectes. Species Général des Lépidop-
téres. V. 7 (Noctuélites, V. 3): 442 pp. Librairie Encyclopédique de Roret,
Paris.

1858. Ibid., 58 Pls. [Noctuélites, 1-24; Deltoides et Pyralites, 1-10;
Uranides, 1; Phalénites, 1-22 and Siculides, 1.]

GunpuacH, J. 1881. Contribucion a la Entomologia Cubana. Pt. 1 Lepiddpteros.
Montiel, Habana. 445 pp.

. 1891. Apuntes para la fauna Puerto-Riquenma. Pt. 7 Lepiddpteros. An.
Soc. Espan. Hist. Nat., 20: 109-207, 323-384.

Harmpacu, F. 1928. A list of the species and descriptions of new forms of the
American genus Zale and a new form of Safia (Lepidoptera: Noctuidae, Cato-
calinae). Trans. Amer. Entomol. Soc., 54: 215-231.

Hampson, G. F. 1898. The Moths of the Lesser Antilles. Trans. Entomol. Soc.
London, 1898, Pt. III, pp. 241-260, Pl. XVII.

. 1913. Catalogue of the Lepidoptera Phalaenae in the British Museum.
V. 13: i-xiv, 1-609, pls. 222-239. Taylor & Francis, London.

HerricH-ScHaAEFFER, G. A. W. 1869. Die Schmetterlinge der Insel Cuba. Noctuina
(part). Corresp.-Blatt zool.-min. Ver. Regensburg, 23(10): 153-160.

Moscuter, H. B. 1890. Die Lepidopteren-fauna von Portorico. Abhandl. Senckenb.
Nat. Ges., 16: 69-360, 1 Pl.

Racugs, P. V. 1914. Museo Cubano “Gundlach” Catalogo General. Zoologia.
156 pp.

Scuaus, W. 1940. Insects of Porto Rico and the Virgin Islands—Moths of the
family Noctuidae. Scientific Survey of Porto Rico and the Virgin Islands,
V. 12, pt. 2, pp. 177-290. New York Academy of Science, New York.

STRAND, E. 1917. Neue Aberrationen der Noctuiden-Subfamilien Hadeninae, Eras-
triinae, Catocalinae, Mominae, und Phytometrinae. Arch. Naturgesch., Abt. A 82,
Heft 2 (1916), pp. 28-50.

VieTtTE, P. 1951. Sur quelques Noctuelles decrites par Guenée (1852-1854). Bull.
Mens. Soc. Linn. Lyon, 20(7): 159-162.

250 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Waker, F. [1858] 1857. List of the specimens of lepidopterous insects in the
collection of the British Museum. Pt. 13, Noctuidae, pp. 983-1236.

1858. Ibid., Pt. 14, Noctuidae, pp. 1237-1519.

Woxcort, G. N. 1923. Insectae Portoricensis (A revised annotated check-list of
the insects of Porto Rico with descriptions of some new species.). Journ. Dept.
Agric. Porto Rico, 7(1): 1-312.

. 1936. Insectae Borinquenses (A revised annotated check-list of the insects

of Puerto Rico.). Journ. Agri. Univ. Puerto Rico, 20(1): 1-600, 190 text figs.

. 1951. The insects of Puerto Rico. Lepidoptera. Journ. Agric. Univ. Puerto

Rico, 32(3) (1948): 537-748, 62 text figs.

Journal of the Lepidopterists’ Society
32(4), 1978, 251-255
A LOST AND MISPLACED TAXON (LEPIDOPTERA:
TORTRICIDAE )

J. F. Gates CLARKE

Department of Entomology, National Museum of Natural History, Smithsonian
Institution, Washington, D.C. 20560

ABSTRACT. The rediscovery of “Antithesia montana” Bartlett-Calvert is recorded.
_ Adult female genitalia are figured, and its assignment to the Tortricidae is established.

In 1893 (p. 831, Pl. 1, Fig. 4) Wm. Bartlett-Calvert described and
figured Antithesia montana from Lolco, Araucania, Chile, without indi-
cation of its family connections. As far as I am able to ascertain, it
was not until 1922 (p. 163) that the species was mentioned again, this
time by Meyrick, when he placed the species in the Oecophoridae, in
the genus Hypercallia. Obviously, Meyrick never saw the species, but
based his placement on misinterpretation of the badly illustrated an-
tennae, as the long labial palpi of Hypercallia.

In January 1974, Dr. Oliver S. Flint, Jr., Department of Entomology,
Smithsonian Institution, rediscovered this species in Argentina, at Pu-
cara, on the Rio Honthue so now it will be possible to establish more
accurately its taxonomic position.

Bartlett-Calvert’s type has disappeared; at least I have not been able
to locate it. Dr. Ariel Camousseight, Chief, Seccion Entomologia, Museo
Nacional de Historia Natural, Santiago informed me that the type is
not in that museum, and Dr. Klaus Sattler, of the British Museum
(Natural History), where some of Bartlett-Calvert’s material was de-
posited, informed me that montana is not represented in that collection.

“Lolco, Araucania” is given as the type locality. “Araucania . . . was
the name of a former region of Chile . . . now mainly comprised in the
provinces of Arauco and Valdivia.” Lolco, however, is now in the
province of Malleco. The specimen before me, collected by Dr. Flint,
came from Pucara, just over the border between Argentina and Chile.
Pucara is situated approximately 225 km south of Lolco and 25 km
west of San Martin de los Andes.

The original description in Spanish is as follows: “Las alas ante-
riores, por encima con la mitad basilar amarilla; la mitad esterna i una
parte de la base negruzca, o con reflejos de luz rojizo-negruzco; en el
centro de la mitad esterna hai una mancha redonda amarilla, encerrada
por un circulo negro; las posteriores de un color moreno-negruzco bril-

bo
Ol
bo

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

2

_ Figs. 1-2. Proeulia montana (Bartlett-Calvert). 1, reproduction of the original
figure; 2, from Argentina, Pucara.

VotuMe 32, NuMBER 4 na 8

liante; por de bajo, todas las alas son morenas inclinandose a negro; la
cabeza de color amarillo; el torax i abdomen moreno oscuro; las franjas
en las alas anteriores son negruzcas i en las posteriores moreno-claras.”

A free translation of the above follows:

“The forewings above from center to base yellow; the outer half and
part of the base dark brown, or with reflections of reddish brown; in
the center of the outer half there is a round yellow spot contained by
a black circle; the hindwings of a shining tawny dark brown; the under-
sides of all the wings are brown inclining to black; the head yellow;
the thorax and abdomen dark brown; the fringes of the forewing are
dark brown and in the hindwings clearly brown.”

The description fits the specimen in hand and needs no emendation.
The female genitalia are described below for the first time. No male
is available.

Genus Proeulia Clarke

Proeulia Clarke, 1962, Proc. Biol. Soc. Washington, 75, 293-294 (Type species.—
Eulia robinsoni Aurivillius, in Skottsberg, The Natural History of Juan Fernandez
and Easter Island, 3: part 2, 266, Pl. 11, fig. 17).

Proeulia montana ( Bartlett-Calvert), new combination

Antithesia montana Bartlett-Calvert, 1893, Santiago de Chile, Univ. Anales, 84:831,
Pl. 1, Fig. 4.

Hypercallia montana (Calvert), Meyrick, 1922, In Wytsman, Genera Insectorum, 180:
163

Male genitalia unknown.

Female genitalia (USNM 24331). Ostium very broad, strongly sclerotized inwardly.
Anirum not differentiated from the strongly sclerotized, very short ductus bursae.
Bursa copulatrix membranous without ventral sclerotized process. Ductus seminalis
from latero-ventral surface of bursa copulatrix.

Type. Lost.

Type locality. Chile, Malleco, Lolco.

Distribution. Chile, Argentina.

Foodplant. Unknown.

Remarks. Although this species lacks one feature characteristic of the genus
Proeulia, the sclerotized process from the ventral surface of the bursa copulatrix, I do
not hesitate to place montana in this genus. As pointed out by Obraztsov (1964),
“Only in the description of the wing venation are some modifications necessary.” He
points out that veins 6 and 7 of the hindwing are sometimes slightly separate, as
opposed to being stalked, as originally described; also that veins 3 and 4 of hindwing
“are either connate or slightly separate at origin” the latter condition found in
montana. Obraztsov also points out that the peculiar process from the ventral surface
of the bursa copulatrix is reduced in some species, and speculates that it might dis-
appear in some taxa. In the case of montana this process is absent, as predicted by
Obraztsov.

Since the identity of this species appears to be beyond doubt, the designation of a
neotype is not necessary.

254 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fig. 3. Proeulia montana (Bartlett-Calvert), ventral view of female genitalia.

VoLuUME 32, NuMBER 4 USS,

ACKNOWLEDGMENTS

The photographs were made by Victor E. Krantz, and the drawing
of the genitalia was made by Mr. George Venable, both on the staff
of the Smithsonian Institution.

LITERATURE CITED

BARTLETT-CALVERT, W. 1893. Nuevos Lepiddépteros de Chile. Santiago de Chile,
Univ. Anales, 84: 813-834, Pls. 1, 2.

Meyrick, E. 1922. In Wytsman, Genera Insectorum. Lepidoptera Heterocera.
Oecophoridae, 180: 1-200, Pl. 1-6.

Osraztsov, N. S. 1964. Neotropical Microlepidoptera, V, Synopsis of the species
of the genus Proeulia from central Chile (Lepidoptera: Tortricidae). Proc. U.S.
Nat. Mus., 116( No. 3501), p. 183-194, Pl. 1-9.

Journal of the Lepidopterists’ Society
32(4), 1978, 256-260

LARISA SUBSOLANA, A NEW GENUS AND SPECIES OF MOTH
FROM EASTERN NORTH AMERICA (OLETHREUTIDAE)

WILLIAM E. MILLER

U.S. Dept. of Agriculture, Forest Service, North Central Forest Experiment Station,
Folwell Ave., St. Paul, Minnesota 55108

ABSTRACT. Larisa Miller, new genus, is proposed for Larisa subsolana Miller,
new species. Larisa is intermediate between the subfamilies Laspeyresiinae and
Eucosminae but is tentatively placed in the former. Larisa subsolana is described
from more than 130 adult specimens representing a geographic range from Texas
and Florida north to Michigan, Ontario, and Massachusetts. Capture dates in Florida
range from March 14 to September 27; elsewhere April 10 to August 7.

I have noticed the species discussed here in museums and private
collections for a decade. It was sometimes identified as Epinotia, Gyp-
sonoma, or Laspeyresia, genera which represent two olethreutid sub-
families. Detailed study eventually showed that the species cannot be
placed in any existing genus. This report is based on more than 130
adult specimens. The letter n denotes number of observations or speci-
mens underlying a particular statement.

Larisa Miller, new genus

Male and female. Head: Maxillary palpus with two developed segments (3 n);
labial palpus slightly upturned, second segment expanding apically; antenna 2/5 xX
forewing length; scaling of front and crown dense, bushy. Thorax: Smooth-scaled;
metathoracic legs unmodified. Forewing: Smooth-scaled; slightly broader toward
termen; costal fold absent; costa slightly and uniformly curved from base to apex;
apex acute; termen convex; dorsum curved; 12 veins, all separate, upper internal
vein of cell arising between veins 10 and 11, vein 11 arising near middle of cell
(Fig. 1,5). Hindwing: Costa convex near middle; apex acute; termen concave;
dorsum straight between veins 1b and 3; pecten normal; veins 3 and 4 stalked to
almost connate; vein 5 straight or slightly bent at base toward 4; veins 6 and 7
stalked (Fig. 1, 5 n). Abdomen: Smooth-scaled; eighth segment of male with a
pair of lateral scale tufts; eighth tergite of female with scales as well as setae.
Male genitalia: Uncus well developed, sclerotized and bifid; gnathos fused ven-
trally across middle; hami long, finger-like; valva simple with rudimentary clasper,
a tuft of fine setae on base of sacculus; one to several long, slender setae may be
present on latero-ventral surface of cucullus; aedeagus sleeve-like, short, tapered;
deciduous cornuti present; dorsal plate of anellus not developed. Female genitalia:
Papillae anales simple; posterior apophyses slightly longer to slightly shorter than
anterior apophyses; sterigma shield-shaped with short finger-like projections beside
ostium, ostium on anterior margin; ductus bursae short, enlarged near middle, sclero-
tized except for a short distance beyond enlargement, convoluted at junction with
corpus bursae; dual thorn-like signa.

Type-species. Larisa subsolana, new species.

Comments. Larisa keys to Laspeyresiinae or Eucosminae (Heinrich 1923,
Obraztsov 1958) depending on character variability and interpretation. It is an
intermediate genus but is tentatively placed in Laspeyresiinae. In the male, the

VoLUME 32, NUMBER 4 ASST

Figs. 1-6. Larisa subsolana, new species. 1, Venation of fore- and hindwing.
2, Fore- and hindwing of specimen from Devil’s Den State Park, Arkansas. Length
of forewing 5.0 mm. 3, Male genitalia of specimen from 3 km E Palmdale, Florida.
4, Enlargement of aedeagus of preceding male. 5, Female genitalia of specimen

from preceding locality. 6, Enlargement of sterigma and associated structures of
preceding female.

958 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

rudimentary clasper and well developed uncus are characteristic of Eucosminae.
Within Laspeyresiinae, Larisa most resembles Laspeyresia and Hemimene or Pammene
(Heinrich 1926, Obraztsov 1960) but differs from both by its convex forewing
termen, long setae on outer surface of cucullus, setal tufts on sacculus, well developed
hami, and in previously enumerated details of forewing or hindwing venation. Larisa
is feminine gender and a patronym for Larisa K. Miller, my volunteer assistant.

Larisa subsolana Miller, new species

Male. Length of forewing 3.8-5.8 mm (71 n). Head: Labial palpus brown,
scales white-tipped, length of second segment 1 x eye diameter and 2.9-4.0 x length
of apical segment as estimated from scaled and descaled specimens (21 n); front
and crown light brown; antenna brown. Thorax: Brown dorsally, including tegula,
scales white-tipped; shining white ventrally; pro- and mesothoracic legs brown on
outer side, scales white-tipped, tarsi white-banded, shining white on inner side;
metathoracic legs shining white. Forewing (Fig. 2): Length 2.6-2.9 x width
(5 n); ground color of upper side brown; basal patch sharply delineated; middle
crossband grayish brown grading apically to darker brown, a thin brown line cen-
trally from costa to dorsum; distal third grayish brown, tinged in costal half with
rust; fringe brown; underside light brown, mottled with white in costal area.
Hindwing (Fig. 2): Widest membranous part 1.1-1.4 x that of forewing (5 n);
upperside, underside, and fringe light brown. Abdomen: Grayish brown dorsally,
paler ventrally, including genital scaling. Genitalia (Figs. 3-4): Width of valval
neck 0.43-0.74 greatest width of cucullus (24 n), the individual values showing
a normal frequency distribution; 8-17 deciduous cornuti or empty cornutus sockets
(2m):

Female. As described for male except forewing length 4.1-6.3 mm (61 n) and
brown genital scaling. Genitalia (Figs. 5-6, 20 n): Sterigma with short finger-like
projections lateral to ostium bursae; posterior apophyses slightly longer to slightly
shorter than anterior apophyses.

Types. HoLtotype ¢: ARKANSAS, Devil’s Den State Park, Washington Co., June
26, 1966 (R. W. Hodges), No. 72093 in National Museum of Natural History.
ALLOTYPE 2: ARKANSAS, same data as holotype except May 30, 1966, in National
Museum of Natural History. Paratypres, 10 specimens: ARKANSAS, same data as
holotype except 22 May 1966, ¢ genitalia slide USNM Tor 2, wing slide WEM 5;
same data as holotype except 20 May 1966, ¢ genitalia slide LKM 1219766; 13 km
SE Ethel, Arkansas Co., 9 July 1969 (R. L. Brown); MISSISSIPPI, Clinton, Hinds
Co., 14 July 1974 (Bryant Mather), No. 73267, 2 genitalia slide LKM 403772;
MICHIGAN, East Lansing, Ingham Co., 15 July 1968 (J. P. Donahue), ¢ genitalia
slide JAB 34; T4N, R2W, Sec. 35, Ingham Co., 12 June 1966 (J. P. Donahue), ¢
genitalia slide KAK 73; ALABAMA, 21 km SW Greensboro, 23 April 1976 (J. B.
Heppner); FLORIDA, 3 km E Palmdale, 4 May 1974 (J. B. Heppner), @ genitalia
slide JBH 455; same data as preceding except ¢@ genitalia slide JBH 454; NEW
YORK, Ithaca, 2 July 1976 (J. G. Franclemont), 2 genitalia slide RLB 645. Para-
types are in National Museum of Natural History; California Insect Survey, Univer-
sity of California, Berkeley; Florida State Collection of Arthropods; Cornell Uni-
versity; University of Minnesota, Twin Cities; and collections of Richard L. Brown,
John B. Heppner, and Bryant Mather. Specimens not designated as paratypes are
in the above repositories, also University of Michigan and Field Museum of Natural
History.

Geographic distribution. Present records for the species occur from Texas and
Florida north to Michigan, Ontario, and Massachusetts (Fig. 7).

Biology. Available biological information is based on adults captured in flight.
The hostplant is unknown. There is probably more than one generation a year.
Capture dates in Florida range from March 14 to September 27 (20 n); elsewhere,

VoLuME 32, NUMBER 4 259

Fig. 7. Distribution of records for Larisa subsolana.

April 10 to August 7 (116 n). If the largest sample of moths from one locality
(46 n, Devil’s Den State Park, Arkansas, 20 May—-22 July 1966, R. W. Hodges)
represents one generation, the flight period is longer than that of many olethreutids
and suggests a protected place of development insulated by shade, soil, or woody
tissue. This sample also shows protandry typical of olethreutids (the median capture
date of males preceding that of females by 12 days) and a male/female ratio of 0.92,
essentially unity.

ACKNOWLEDGMENTS

I thank the following for specimen loans and other assistance: Jerry
A. Powell, University of California, Berkeley; Don R. Davis, National
Museum of Natural History; Roland L. Fischer, Michigan State Uni-
versity; Thomas E. Moore, University of Michigan; Bryant Mather,
Clinton, Mississippi; Charles P. Kimball, West Barnstable, Massachu-
setts; and Henry Dybas, Field Museum of Natural History. I am espe-

260 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

cially indebted to Richard L. Brown, Cornell University, and John B.
Heppner, University of Florida. After more than 50 Larisa specimens
had come to my attention, I prepared a draft of this paper and asked
several workers including Brown and Heppner to review it. Unknown
to me, both were also independently studying the insect and each
generously forwarded research information and more than 60 additional
specimens to me.

LITERATURE CITED

Hernricu, C. 1923. Revision of the North American moths of the subfamily
Eucosminae of the family Olethreutidae. U.S. Natl. Mus. Bull. No. 123, 298 pp.

1926. Revision of the North American moths of the subfamilies Laspey-
resiinae and Olethreutinae. U.S. Natl. Mus. Bull. No. 132, 216 pp.

Opraztsov, N. 1958. Die Gattungen der Palaearktischen Tortricidae. II. Die
Unterfamilie Olethreutinae. Tijdsch. Entom. 101: 229-261.

. 1960. Die Gattungen der Palaearktischen Tortricidae. II. Die Unter-

familie Olethreutinae. 3. Teil. Tijdsch. Entomol. 103: 111-143.

Journal of the Lepidopterists’ Society
32(4), 1978, 261-272

NOTES ON MEXICAN ACTINOTE (NYMPHALIDAE: ACRAEINAE)
AND THEIR RELATIVES, WITH DESCRIPTION OF
A NEW SUBSPECIES

JACQUELINE Y. MILLER AND LEE D. MILLER
Allyn Museum of Entomology, 3701 Bay Shore Road, Sarasota, Florida 33580

ABSTRACT. The application of names described for South American species to
their Central American, and especially Mexican, counterparts has led to great con-
fusion in the literature. Actinote stratonice oaxaca is described from Oaxaca, Mexico;
this insect had been reported previously as the nominate subspecies. Three species
of the Actinote thalia group, A. calderoni, lapitha and thalia are illustrated, redescribed
and discussed. A key for the separation of the three species is provided. The first
two species are recorded from Mexico—calderoni had been previously misidentified
as lapitha from there. One name, A. lapitha zilchi Franz and Schroder, is synony-
mized to calderoni.

Collecting in Mexico over the past forty years has yielded many
butterfly species that were previously unknown from there, not a few
of which were totally unexpected. These unexpected taxa have created
many problems, usually for one of two reasons: 1) the butterfly was
an already described Central or South American species and was de-
scribed as new from Mexico because of a lack of comparative material
or an ignorance of the pertinent literature; or 2) many species (espe-
cially those figured and described in Seitz) incorrectly have been as-
cribed to the Mexican fauna, again because of a lack of adequate
comparative material. Both situations are well demonstrated in the
Nymphalidae: Acraeinae. The Hoffmann (1940) catalog lists only four
species of this subfamily within the borders of Mexico, but recent
collecting has uncovered one that has been misidentified in collections
and in correspondence—the one with which the first-mentioned species
had been confused and an undescribed subspecies of a well-known
South American insect. In the hope of unravelling the confusion in
this small subfamily (within the Mexican borders), we offer these
notes.

Actinote stratonice oaxaca J. Miller and L. Miller, new subspecies
Figs. 1-6

Male. Head, thorax and appendages black; abdomen black with a reddish-brown
midsternal line. Upper surface of wings similar to that of the nominate subspecies,
but paler, and with the following differences: forewing totally black anteriad of cell
(partially reddened in other subspecies ); dark marking at end of forewing cell much
smaller than in other subspecies and black area at base of forewing cell and along
inner margin more restricted than in other populations. Under surface pattern
also paler than in s. stratonice with forewing differences as noted for upper surface

bo
o>)
bo

JouRNAL OF THE LEPIDOPTERISTS SOCIETY

Figs. 1-4. Actinote stratonice oaxaca J. Miller and L. Miller, new subspecies.
1-2, Holotype ¢, upper (1) and under (2) surfaces; MEXICO: OAXACA: Can-
delaria Loxicha, 550 m. 3-4, Paratype 2, upper (3) and under (4) surfaces; same
locality. Both specimens in Allyn Museum of Entomology.

and in addition the pale orange patches in the forewing cell area darkened basad,
blending to tawny-yellow distad; black areas admixed with pale yellow scales, espe-
cially in base hindwing. ¢ genitalia as illustrated, differing from those of other
stratonice in only minor respects. Length of forewing of Holotype ¢ 24.4 mm,
those of eleven of the @ Paratypes range from 26 to 31 mm.

Female. Differs from the 2 of nominate stratonice in the same manner as does
the @, but additionally the basal black areas of the forewing encompass the proximal
third of the cell and below it along the inner margin, and all of the black areas
below are completely suffused with yellow scaling. One @ Paratype has a hindwing
supernumerary vein off Rs on the right side. @ genitalia as illustrated and com-
paring well with those of other subspecies. Lengths of forewings of 12 2 Para-
types range from 28.2 to 38 mm.

Specimens examined. Described from 25 specimens, 13 males and 12 females,
from the state of Oaxaca, Mexico.

Types. Hotorype ¢: MEXICO: OAXACA: Candelaria Loxicha, 550 m, 8.ix.
1969 (KE. C. Welling). Pararypres: all MEXICO: OAXACA: same locality as
Holotype, 14 15.xi.1967, 16 15.ix.1968, 12 21.vii.1970, 14 19 27.viii.1970, 19
21.vii.l1973 (all E. C. Welling M.); El Portillo del Rayo, Candelaria Loxicha, 14
32 17.xi.1967 (all E. C. Welling M.), 64 49 18.vii.1976 (all de la Maza family);
Rio Molina, Mpio. Suchistepec, 2200 m., 19 10.x.1967 (E. C. Welling M.); San
Jose Pacifico, Mpio. Rio Hondo, 2400 m., 14 9.x.1967 (E. C. Welling M.); Puente

VoLUME 32, NuMBER 4 263

Figs. 5-6. Actinote stratonice oaxaca J. Miller and L. Miller, new subspecies.
5, 6 genitalia of Paratype; MEXICO: OAXACA: Candelaria Loxicha; preparation
M-3630 (Jacqueline Y. Miller). 6, 2 genitalia of Paratype; same locality; prepara-
tion M-3602 (Jacqueline Y. Miller).

del Guajolote, Jalatengi, 19 21.iii.1975 (E. Fernandez), 14 21.xi.1975 (de la Maza
family); Dos de Mayo, 1¢ 18.v.1976 (de la Maza family).

Disposition of type series: Holotype ¢, two ¢ and four @ Paratypes in the
collection of the Allyn Museum of Entomology; two ¢ and three @ Paratypes in
the collection of E. C. Welling M. and eight ¢ and five 2 Paratypes in the de la
Maza collection.

Remarks. The subspecific name refers to the state from whence the new sub-
species came.

A. stratonice is recorded in Seitz from the Sierra Madre de Santa
Marta of Colombia and southward through the mountains of Venezuela
and into Ecuador. The present subspecies was first recorded in the
literature from Mexico by de la Maza R. and de la Maza E. (1975),
but at the time they did not recognize it as a separate subspecies.

The subspecies oaxaca is characterized by its overall dull coloration
and by the suffusion of the under surface with pale yellow scales. In
none of the other subspecies of stratonice do these characters appear.
The disjunct distribution of stratonice, with a Mexican subspecies widely
separated from its nearest relatives, is indeed intriguing.

264 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Figs. 7-10. Actinote calderoni Schaus. 7-8, 4, upper (7) and under (8) sur-
faces; MEXICO: CHIAPAS: Mapastepec. 9-10, 2, upper (9) and under (10)

surfaces; same locality. Both specimens in Allyn Museum of Entomology.

The Identity of the Mexican Actinote thalia Group Species

Pale Mexican specimens of thalia group Actinote in most collections
usually have been identified as A. lapitha (Staudinger). Comparison
of most Mexican material with Staudinger’s description and subsequent
literature citations suggest that these specimens could not be true
lapitha. Accordingly, we searched museum and private collections and
the literature for documented material that might shed light on the
correct name for the Mexican insects. The results were surprising, and
to avoid future confusion the following redescriptions are offered to
aid in the identification of these butterflies.

Actinote calderoni Schaus, 1920
Figs. 7-12

Actinote calderoni Schaus, 1920: 434 (TL—Anteos, El Salvador).
= Actinote lapitha zilchi Franz and Schréder, 1954: 80; fig. 2 (TL—Km. 30, Son-
sonate Rd., La Libertad, El] Salvador). [New Synonymy. ]

Male. Head, thorax and first two abdominal segments dark brown covered
with a few fuscous and tawny dorsal hairs; remaining abdominal segments naked

VoLUME 32, NUMBER 4 265

Figs. 11-12. Actinote calderoni Schaus. 11, @ genitalia; MEXICO: CHIAPAS:
Mapastepec; preparation M-3633 (Jacqueline Y. Miller). 12, 2 sterigma; same lo-
cality; preparation M-3601 (Jacqueline Y. Miller).

dorsad; thorax and first two abdominal segments covered with a few tawny ventral
scales; last abdominal segments ventrally naked; pleural line buff. Palpi tawny with
a few black scales along inner surface only. Antennae and legs black.

Upper surface of wings very thinly scaled with white, giving a dusky, off-white
appearance; forewing smoky at apex and marginally to Cuz-2A; veins slightly dark-
ened and interneural spaces with single smoky stripes. Under surface of wings
similar, but a faint, dark comma-shaped marking lies from M;-Cu: to Cui-Cuz of the
forewing and numerous long, dark hairs lie on the veins, especially of the hindwing.
6 genitalia as illustrated. Forewing lengths of the ¢ examples examined range
from 21.5 to 24.5 mm.

Female. Similar to ¢, but paler, especially the forewing smoky markings above
and below. @ genitalia as illustrated, generally characterized by having the sterigma
more sculptured than in related species. Lengths of forewings of 2 specimens at
hand range from 25 to 30.5 mm.

Types and specimens examined. We have examined 15 specimens, seven males
and eight females, from the following localities: EL SALVADOR: Anteos, 1¢
(Hotoryee, USNM), 12 (possible Paratype, CM). GUATEMALA: Tiquisate,
28.vi.1947, 12 (AMNH). MEXICO: CHIAPAS: Mapastepec, various dates, 1939-
1959, 24 59 (AME), 3é6 12 (AMNH); San Jeronimo, 600 m., 17.vii.1975, 192
(E. C. Welling M.).

The records from Mexico and Guatemala are apparently the first
for either country. The present insect has been masquerading in Mex-
ican collections for years as A. lapitha (Staudinger), a species that is
abundantly distinct.

Not only have workers on Mexican butterflies been confused on the
identity of calderoni, but also those in other parts of Central America.
The fact that Schaus’ (1920) description appeared in an entomologically
obscure journal has resulted in the paper never being cited previously
by workers on Actinote. Were the “Fifty-Year Rule” still in effect in

266 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Figs. 13-16. Actinote lapitha (Staudinger). 13-14, ¢, upper (13) and under
(14) surfaces; MEXICO: CHIAPAS: Tierra Blanca, Mpio. La Trinitaria, 1500 m.
(E. C. Welling M. collection). 15-16, 2, upper (15) and under (16) surfaces; no
locality data (AMNH collection ).

the International Code of Zoological Nomenclature, the name calderoni
could be ignored with impunity, but happily this rule was repealed
a few years ago, so we must return to the oldest name. Franz and
Schroder (1954) had obviously not seen the Schaus description, but
their excellent figure of the type of A. lapitha zilchi is referable to
specimens of calderoni that we have seen (including the Holotype),
and since the two taxa were described from within 100 km. of one
another, it is evident that they represent the same species.

Actinote lapitha (Staudinger), 1888
Figs. 13-18

Acraea lapitha Staudinger, 1888: 82 (TL—“Chiriqui” ).

Male. Head, thorax and abdomen clothed with black dorsal hairs; head, palpi
and most of thorax also clothed with black ventral hairs; small patch of tawny scales
on meso- and metathoracic preepisterna and an additional such patch on metathoracic
epimeron; abdomen clothed with admixed fuscous, tawny and buff scales; pleural line
tawny and buff only. Antennae and legs black. Ground color of forewing above
translucent and tawny with margins outlined in dull gray-brown, especially at apex;
prominent gray-brown transverse marking from end cell to Cu.-2A, interspersed with

VoLUME 32, NuMBER 4 267

Figs. 17-18. Actinote lapitha (Staudinger). 17, ¢ genitalia; MEXICO: CHI-
APAS: Tierra Blanca, Mpio. La Trinitaria; preparation M-3416 (Jacqueline Y. Mil-
ler). 18, 2 sterigma; no data; preparation M-1657 (Jacqueline Y. Miller).

tawny scaling from end cell to M;-Cu:; veins and interneural striping prominent,
gray-brown. Hindwing above also translucent tawny, dull gray-brown at costa and
along margin; veins heavily darkened with gray-brown and interneural markings of
same color short and heavy. Forewing below similar to upper surface, but margins
and apex not so dark and with an additional dull gray-brown bar across middle of
cell. Hindwing below as above with a faint gray-brown cell end marking from
costa to base of M; and more prominent veinal and interneural blackish-brown striping.
Fringes of both wings blackish on both surfaces. ¢ genitalia as illustrated. Lengths
of forewings of the three ¢ specimens examined range from 22.8 to 23.6 mm.

Female: Similar to ¢, but thorax sparsely clothed with tawny scales, markings
of all wings paler, but extra-discal band of forewing more prominent and base of cell
of same wing somewhat overscaled with fulvous. 2 genitalia as illustrated; sterigma
somewhat heavier than that of the next species and not quite so ornamented as in
calderoni. Lengths of forewings of the two 2 examples before us 20.2 and 23.1 mm.

Types and specimens examined. We have seen two females and three males of
this insect. PANAMA: Jicaron Island, 14—15.i.1902, 1¢ (BMNH). COSTA RICA:
Puerto Golfito, 4.vii.1965, 1¢ (Gordon B. Small, Jr. collection). MEXICO: CHI-
APAS: Tierra Blanca, Mpio. La Trinitaria, 1500 m., 15.ix.1972, 1¢ (E. C. Welling
M.) No Data, 22 (AMNH).

Evidently the Costa Rican record is a new, but not unexpected one.
Hoffmann (1940: 672) lists lapitha from “Tierra caliente de la costa del
Pacifico de Chiapas,” no true specimens of that species are in the Hoff-
mann collection in the AMNH. All of the specimens in Hoffmann’s
material were calderoni, and one of these bore a determination label
in Hoffmann’s hand of “Actinote lapitha Staudinger.” Since Hoffmann
obviously confused lapitha with calderoni, we feel that Mr. Welling’s
specimen of the former is the first authentic record from Mexico.

A. lapitha was described from the Chiriqui region of Panama, but
the type specimen is apparently no longer extant, perhaps having been

68 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

Figs. 19-22. Actinote subhyalina (Staudinger). 19-20, ¢, upper (19) and under
(20) surfaces; PERU: Rio Cachiyacu, Iquitos (BMNH). 21-22, Lectotype @ (see
designation in text), upper (21) and under (22) surfaces; PERU: Yurimaguas
(ZMHU).

destroyed during World War II (H. J. Hannemann, pers. comm.). We
have been unable to locate an authentic Chiriqui specimen in more
recent collections; hence, we do not designate a Lectotype here.

Actinote subhyalina (Staudinger), 1888
Figs. 19-24

Acraea subhyalina Staudinger, 1888: 81; pl. 32 (TL—Yurimaguas, Peru).

Description. Sexes similar; Head, thorax and abdomen above and below
blackish-brown clothed with fuscous to buff scales. Palpi blackish-brown; legs
brown and antennae dull reddish-brown. Forewing above buff, translucent and
laved with fulvous toward base (darkest in @); apex and margin dull brown; veins
darkened; transverse markings: a jagged blackish-brown one across cell about %4
distance from base to end and a second one from costa across end cell to anal margin
(not prominent below Cu; in ¢). Hindwing above buff, translucent, with darker
veins; margin outlined in dull brown; interneural striping in anal region; prominent
dark brown bar from costa to Cu;-Cuz and a dull brown mark across distal end of
cell. Under surface of wings as above, but dull brown markings slightly overscaled
with buff, hindwing distal band more diffuse and an additional dull brown stripe in
hindwing cell Sc+Ri-Rs. Lengths of forewings of all specimens examined ranged
from 20 to 22 mm. Genitalia of ¢ and 9° as illustrated.

Specimens examined. We have been able to examine only a single ¢ and six
? specimens. PERU: Rio Cachiyacu, Iquitos, [18]93, Stuart, 14 39 (BMNH);
Yurimaguas, 19 (ZMHU); No data, 22 (CM).

VoLUME 32, NUMBER 4 969

Figs. 23-24. Actinote subhyalina (Staudinger). 23, 4 genitalia; PERU: Rio
Cachiyacu, Iquitos; preparation M-3403 (Jacqueline Y. Miller). 24, @ genitalia of
Lectotype; PERU: Yurimaguas; preparation M-3411 (Jacqueline Y. Miller).

Staudinger (1886: 81) described A. subhyalina from 12 2 specimens
taken at Yurimaguas. Certainly the figured 2° from Berlin was one of
Staudinger’s syntypes, and the two specimens from CM may have been,
but in the case of the latter two specimens this cannot be ascertained
with precision. One of the CM specimens bears the label “Acraea
subhyalina/ from Dr. O. Staudinger/1885,” a date that was three years
before the description; the second specimen bears a number (Stauding-
ers?) only, “386.” The specimen received from the ZMHU is defi-
nitely from the type locality, was in the Staudinger collection and has
been labelled as “Origen” by Staudinger or someone subsequent to him.
It is the logical candidate for designation as the Lectotype of the name,
and we have so labelled it, affixing a red, partially printed, partially
handwritten (italics) label to it: “Lectotype/ Acraea/ subhyalina/
Staudinger, 1888/ designated by Jacqueline Y. Miller/ & Lee D. Miller,
HOLT.

DIscUusSION

The impetus for this project was a series of seven specimens in the
Allyn Museum collection from Mapastepec, Chiapas, Mexico. Exami-
nation of these specimens revealed that while they were closely related
to A. lapitha (the name associated with them), they were abundantly
distinct.

Letters for additional specimens brought two from the AMNH that
were in agreement with the original description of lapitha (but without
data) and five more of the odd one, four of which were from the

270 JoURNAL OF THE LEPIDOPTERISTS SOCIETY

Hoffmann collection, taken by Dr. Escalante. A later trip to CM yielded
one more specimen and the first clue to the identity of the Mexican
material that we had. The additional label on the CM specimen stated
that it was “Actinote calderoni Schaus,” but at the time we were unable
to find the original description or any reference to it, and we still as-
sumed that the name might have been a manuscript name only. Re-
checking the series and the type collection at the USNM yielded not
only the type specimen of calderoni, but also finally the reference to
the original description of this elusive name.

The confusion did not end there, though. The two true lapitha from
the AMNH had a fulvous basal flush on the forewing, thus resembling
subhyalina. This led us to wonder if the specimen figured by Staudinger
(1888) was subhyalina or lapitha. Letters to the BMNH and the ZMHU
brought additional specimens of Staudinger’s insects and some very
helpful information.

All three species are rare in collections, especially the males. Since
we encountered such difficulty in making determinations in the thalia
group, we present the following key to aid other workers to the species
treated here.

1. Forewing above with dark transverse band at end cell from costa to near
anal angle: 2-222. Sone nce ee 2.
1’. Forewing with no such band; Mexico to El Salvador calderoni Schaus.
2. Hindwing above with prominent dark discal markings at end of cell; Peru
PE pk lial RE 8 cca are ed TEE Ly ail al Shah subhyalina (Staudinger).
2’. Hindwing above without prominent marking at end cell; Mexico to Panama
eg dont GM Nu | aetna Di eth A es lapitha (Staudinger).

Essentially the key characters for the separation of the species in this
complex are the dark bars across the cells of both wings. A. subhyalina
shows these bars on the upper surfaces of both wings, A. lapitha has
only the one on the forewing and A. calderoni has neither. Seemingly
the orange flush at the base of the upper forewing should be diagnostic,
but whereas it is prominent in most subhyalina, it also appears in some
female lapitha, hence it is diagnostic of neither. |

The male genitalia (Figs. 11, 17, 23) are similar, but subtly different,
in all three species. The valvae are elongated and slightly curved dorsad
in calderoni, whereas they are squarecut posteriad in both lapitha and
subhyalina. The saccus of subhyalina is much more elongate than is
that of either of the other two species.

Most of the differences between females of these species lie in the
sterigmal region. The posterior margin of the lamella postvaginalis is
U-shaped and narrow in lapitha, U-shaped and expanded in subhyalina
and even more enlarged and W -shaped in calderoni. The entire opening

VoLUME 32, NuMBER 4 TEAL

of the ostium bursae is darkly sclerotized in a narrow ring in lapitha,
a somewhat broader darkly sclerotized ring in subhyalina but only
darkly sclerotized in four separate areas around the opening in calderoni,
but the area around the ostium itself is lightly sclerotized. The genital
capsule is larger in calderoni than in the other species.

The tarsi, as is true of all members of the thalia group, are asym-
metrical and comparable one with another.

Geographically A. subhyalina can be immediately separated from
lapitha and calderoni, none of the three species being found in Colombia
or Ecuador, as far as we know. It would not be surprising to see
subhyalina from at least the latter country, and there might be sym-
patry between subhyalina and lapitha in Colombia. A. lapitha, as re-
corded here, has a much more extensive range than previously believed,
and calderoni is not restricted to El Salvador. Either of these species,
or both, may well be found in Honduras and Nicaragua. We hope
that this paper will encourage others to try to fill in the distributional
blanks for this interesting group.

ACKNOWLEDGMENTS

We are deeply indebted to the following for loan of material and for
access to their collections (abbreviations) of Actinote: Dr. F. H. Rindge,
American Museum of Natural History (AMNH); Mr. H. K. Clench,
Carnegie Museum of Natural History (CM); Mr. W. D. Field, National
Museum of Natural History (USNM); Mr. P. R. Ackery, British Museum
(Natural History) (BMNH); Dr. H. J. Hannemann, Zoologische Museum
der Humboldt Universitat (ZMHU); Dr. T. Escalante and Messrs. R.
de la Maza R. and J. de la Maza E., Mexico, D. F., Mexico; Mr. E. C.
Welling M., Merida, Yucatan, Mexico and Mr. G. B. Small, Jr., Balboa,
Canal Zone. The Allyn Museum of Entomology is abbreviated AME
in part of the paper.

Mr. Field and Dr. J. F. G. Clarke of the USNM were instrumental
in obtaining a copy of the Schaus paper for us. We also thank Mr.
S. R. Steinhauser for pointing out the Franz and Schroder paper to us
and to Mr. H. W. Dybas of the Field Museum for obtaining a copy
of it for our use.

Special thanks are due Mr. A. C. Allyn of this institution for taking
the photographs contained herein and for reading the manuscript.

LITERATURE CITED

Franz, E. & H. Scurover. 1954. Tagfalter (Lep. Rhopalocera) aus El Salvador.
Senckenbergische Biol. 35: 75-87; figs.

bo
~l
bo

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

HorrMAnn, C. C. 1940. Catalogo sistematico y zoogeografica de los Lepidopteros
mexicanos. la. Parte. Papilionoidea. An. Inst. Biol. 11: 639-739.

Jorpan, K. 1924. Acraeidae. In Seitz, A., Macrolepidoptera of the World 5: 358—
374.

DE LA Maza R., R. & J. DE LA Maza E. 1975. Adiciones al Catalogo de Lepidop-
teros mexicanos. Rev. Soc. Mexicana Lepid. 1: 62-63; figs.

Scuaus, W. 1920. Descriptions of two new species of butterflies from tropical
America. J. Washington Acad. Sci. 10: 434-435.

STAUDINGER, O. 1888. In Staudinger, O. and E. Schatz, Exot. Schmett. (1): 81-
oo. Pio:

Journal of the Lepidopterists’ Society
32(4), 1978, 273-276

PAPILIO ARISTODEMUS (PAPILIONIDAE) IN THE BAHAMAS

Harry K. CLENCH
Carnegie Museum of Natural History, Pittsburgh, Pennsylvania 15213

ABSTRACT. Two subspecies of Papilio aristodemus Esper, both new, are de-
scribed from the Bahamas: driophilus (TL: Cutlass Bay, near Dolphin Head, Cat
Island), known from Cat, South Andros, and North Andros islands; and bjorndalae
(TL: Man of War Bay, Great Inagua Island), known only from Inagua and strikingly
different from any known subspecies, though apparently derived from driophilus.

Papilio aristodemus is an Antillean swallowtail with a strong tendency
to vary geographically. Nominate aristodemus Esper 1794 occurs on
Hispaniola; the subspecies temenes Godart 1819 is found on Cuba and
on Little Cayman in the Cayman Islands (Carpenter & Lewis 1943);
subspecies ponceanus Schaus 1911 is known only from southeastern
Florida, particularly Key Largo. An old record of the species for Puerto
Rico (cf. Comstock 1944: 535), subspecies unknown, is not substantiated
by more recent captures.

This species recently has been discovered in the Bahamas (Clench,
1977). I first found it on South Andros Island in early June 1974. A
year later, in early June 1975, I took it also at the southern end of
Cat Island, and in 1976 I collected a specimen on North Andros. The
populations on these islands are not absolutely identical, but they are
close enough to be referred to the same subspecies, described below as
driophilus.

Miss Karen Bjorndal, a graduate student at the University of Florida,
Gainesville, spent over a year on Great Inagua Island, from April 1975
to August 1976, studying the energy budget and nutritional ecology of
the Green Turtle, Chelonia midas. While there she also made a collec-
tion of butterflies, which she has generously donated to Carnegie Mu-
seum of Natural History. In her collection are two specimens of a
striking new subspecies of aristodemus, in several ways the most distinct
of all. It is a pleasure to name it in honor of Miss Bjorndal.

Papilio aristodemus driophilus, new subspecies

Papilio aristodemus ponceanus: Clench 1977:190.

Description. Much closer to ponceanus (Florida) than to either a. aristodemus
(Hispaniola) or temenes (Cuba). This is shown particularly by its sharing with
ponceanus such traits as the thin median yellow-ocher band, and the complete, only
slightly curved, subterminal row of yellow-ocher lunules, both on the forewing above.
From ponceanus, however, it differs in these ways:

(1) On the hindwing upperside the subterminal yellow, orange, or red-orange spot

274 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

in Cu-2A is completely separated by a black bar from the terminal yellow or orange
distad. In ponceanus these two pale areas are connected by a narrow isthmus along
Cuz.

(2) On the forewing above, the cell is rather densely and evenly sprinkled with
pale (greenish) scales. In ponceanus the sprinkling is extremely sparse and tends to
be limited to the basal and costal parts of the cell.

(3) The projections of the hindwing termen at the vein-ends (including the tail)
are longer than they usually are in ponceanus (but the latter is inclined to be variable
in this respect).

(4) On the forewing above, the segment of the median band in M:-Mb is broadly
in contact with the next posterior segment, in M2-M;. In ponceanus the M:-M2 seg-
ment is smaller, and posteriorly separated from the next one by a fuscous gap or
(rarely) touches the next segment at a point only: the two segments are never in
broad contact.

(5) On the hindwing upperside the median pale band is somewhat broader than
in ponceanus. In driophilus the segment in the cell is consistently wider than the
fuscous in the cell just distad; in ponceanus the pale band is here subequal to the
fuscous in width, or it is somewhat narrower.

Length of forewing. Male, one only, 47.0 mm; female, 48.0-50.0 mm, mean
(of 4), 49.1 mm. Measurements are of the type series only.

Types. Ho.otype, 2, Cutlass Bay, near Dolphin Head, southern Cat Island,
Bahamas, 6.vi.1975, leg. H. Clench, sta. 259 b; C. M. Acc. 27783. Paratypess, 13
3@, as follows: 22, same data as holotype; 1¢ 12, the same except 4.vi, sta. 257 b.
Holotype and paratypes, C. M. Ent. type series no. 680.

Remarks. In addition to the type series I have examined 4¢ 22 from South
Andros Island, Bahamas: ca. 2 mi S Driggs Hill, 2-8.vi.1974, leg. H. Clench. The
forewing length of this series is as follows: males, 43.5-49.0 mm, mean (of 4),
45.0 mm; females, 47.5--51.0 mm, mean (of 2), 49.2 mm. These specimens agree
closely enough with the Cat Island series, notably in all points mentioned in the
above description, that I believe them correctly referred to the subspecies driophilus.
The agreement, however, is not perfect and the two island samples show a few,
mostly statistical, differences:

(a) On the hindwing above, the subtornal pale spot (as in (1) above) is dark
orange, with little or no pale edging, in all the Cat Island specimens; it is light
orange, more or less heavily edged laterally with yellow, in all but 1¢ from South
Andros (in which it is dark orange). (In ponceanus; dark orange with slight lateral
yellow. )

(b) On the hindwing above, a small rusty spot in the base of cell M3-Cu: is present
in 12 12 (40%) of the Cat Island series, but is totally absent from the South
Andros series. (In ponceanus: 56%).

(c) The discal cell on the forewing underside is filled with smooth, pale yellow-
ocher in all Cat Island specimens; in all South Andros specimens the cell has periph-
eral fuscous and faint distal longitudinal fuscous streaks. (In ponceanus: as on
South Andros, but the fuscous is even heavier. )

(d) On the forewing underside, the subapical transverse fuscous bar from costa
(just distad of, and parallel to, the conspicuous pale bar on the forewing upperside )
extends inward to cross cell Rs-M: in 1¢ (17%) from South Andros, in 42 (80%)
from Cat Island. In the remaining individuals it does not reach that interspace (In
ponceanus: 22%.)

On South Andros driophilus flew in dense scrub, usually 1-2 m above
the ground, only briefly and occasionally pausing to feed at the flowers

of shrubs in that height range. The butterflies were mostly in the scrub
itself and they entered roadways or other open areas only to cross from

VoLUME 32, NUMBER 4 UTS

one part of the scrub to another. These habits they shared fully with
P. andraemon bonhotei Sharpe 1900, which flew with driophilus, and
the two were virtually indistinguishable on the wing. On Cat Island
the habits of driophilus were similar except that individuals were seen
more often in open areas, especially at the flowers of ornamental vines
and shrubs around the hotel where I stayed.

On 28 September 1976 I took a single male driophilus just north of
Nicolls Town, North Andros, a new record for that island. I saw no
others and am at a loss to explain the late capture date. The butterfly
is quite fresh and was found flying in a somewhat overgrown old field.
It, too, is referable to the new subspecies, although differing in a few
respects (e.g., the median pale band on the hindwing above is thicker
than in any other driophilus seen except one of the female paratypes
from Cat Island; and its distal edge is straight [as in bjorndalae], not
convex near Rs and M,). With regard to traits (a) through (d) above:
(a) the subtornal pale spot is dark orange with slight lateral yellow
(as in ponceanus); (b) it has no rusty spot in M,;—-Cu,; (c) on the
forewing below it agrees with South Andros specimens in the discal
cell coloration; (d) also on the forewing below, the subapical fuscous
bar extends inward only to R;.

Papilio aristodemus bjorndalae, new subspecies

Description. Differs in two major traits from all previously known subspecies of
aristodemus: (1) a large patch of rusty red is present on the hindwing upperside
between M: and the inner margin, and between the cell-end and the diffuse, faint
band of sprinkled blue scaling that basally edges the subterminal row of pale spots;
and (2) on the hindwing, both above and below, the subterminal pale spots posterior
to Mz are distally displaced and reduced in size, so that the row is essentially
parallel to the termen throughout and the component spots are of similar thickness
and more quadrate (less lunular). The latter trait is particularly conspicuous on
the underside. The rusty red patch varies in the two specimens at hand, but I can-
not tell whether the variation is sexual or individual. In the female the patch is
large, the component spots contiguous, and there is even a minute extra dot of the
same color in Mi-M2; in the male the component spot in Cui-Cu: is wanting, and
those in Me-M;-Cu; are thin and short, separated by fuscous along the veins.

The median pale band of the forewing upperside is thin, as in subspecies pon-
ceanus, driophilus, and aristodemus, and slightly or not at all broken at Mb, as in
driophilus; on the forewing upperside the subterminal row of pale spots is lightly
curved (as in all subspecies except nominate aristodemus, in which it is strongly
curved, almost angulate, near Cui), and continues strongly costad to Ri, as in pon-
ceanus (in a. aristodemus it stops at M2; in temenes at about M,; in driophilus at
R; or Ru, the segment in R.-R;s being often weak or wanting). On the hindwing
upperside, in Cu-inner margin, the subterminal pale bar is connected to the pale
terminal area by a narrow isthmus along Cu, as in ponceanus (in all other sub-
species the two pale areas are usually completely separated by intervening fuscous).
The median pale band on the hindwing upperside is thin, about as in ponceanus
or even thinner, and about half as thick as that in driophilus, and its distal edge is
straight, not convex near Rs, as it is in ponceanus and driophilus. This median band

276 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

posteriorly curves distad at the inner margin and runs along the margin almost to
the blue bar, as in driophilus (in the others it intersects the inner margin at a high
—often right—angle and does not run distad). Basad of the subterminal pale spots
is a band of sprinkled blue scales, strongest in Cu:-2A but extending, weaker, costad
to M, or Rs, essentially as in driophilus and ponceanus (in nominate aristodemus
it is absent except for the segment in Cuz-2A, and in temenes it is usually so).
Both specimens are smaller than any other aristodemus I have seen.

Length of forewing. Male, 40.0 mm; female, 44.0 mm.

Types. Ho.orypee, 2, Man of War Bay, Great Inagua Island, Bahamas, 4.x.1975,
leg. Karen Bjorndal. C. M. Acc. 29104. Paratype, ¢, Calf Pond, northwestern
Great Inagua, 18.v.1976, leg. Karen Bjorndal. C. M. Acc. 29104. Holotype and
paratype, C. M. Ent. type series no. 690.

Remarks. This subspecies apparently was derived from driophilus of the central
Bahamas, although it has departed from it to an unusual and striking degree. The
large rusty red patch on the hindwing above gives it a distinctive appearance, but
the patch is foreshadowed by the small, obscure, rusty red spot that appears in
M;-Cu; in some driophilus and ponceanus (see character (b) in the Remarks under
driophilus above).

Miss Bjorndal comments (in litt.): “From September to December
[1975] and from May to August [1976] swallowtails were flying on
Inagua. I was unable to distinguish which species [aristodemus or
andraemon|. They were commonly seen in open scrub, dense scrub,
coppice, coastal areas and [in the residential area of | Matthew Town.”

In April 1977 on Little Inagua Island I repeatedly saw, but was
unable to capture, a swallowtail in the short, narrow strip of low forest
on the western coast, about a mile south of Northwest Point. Like Miss
Bjorndal, I was unable to tell which of the two species it might have
been.

LITERATURE CITED

Carpenter, G. D. H., & C. B. Lewis. 1943. A collection of Lepidoptera (Rho-
palocera) from the Cayman Islands. Ann. Carnegie Mus. 29: 371-396, ill.
Crencu, H. K. 1977. A list of the butterflies of Andros, Bahamas. Ann. Carnegie
Mus. 46: 173-194, ill.

Comstock, W. P. 1944. Insects of Porto Rico and the Virgin Islands, Rhopalocera
or butterflies. Scient. Survey Porto Rico and the Virgin Islands (New York
Acad, Sci.) 12(4): 421-622, ill.

Journal of the Lepidopterists’ Society
32(4), 1978, 277-281

THE NAMES OF CERTAIN HOLARCTIC HAIRSTREAK GENERA
(LYCAENIDAE)

Harry K. CLENCH
Carnegie Museum of Natural History, Pittsburgh, Pennsylvania 15213

ABSTRACT. The palearctic genus Strymonidia Tutt 1908 and the nearctic genus
Euristr'ymon Clench 1961, both in current use, are both synonymized to Fixsenia
Tutt 1907, on the basis of male genital structure. The currently used palearctic
genus Nordmannia Tutt 1907, and many other generic names not in general use,
must be synonymized to Satyrium Scudder 1876, on similar structural evidence. Both
Fixsenia and Satyrium are shown to be holarctic, and both are unusually variable in
external facies. All species known to belong to both genera are listed.

The correct generic names to be applied to some of the palearctic
hairstreaks has long been a problem. The species concerned are those
listed by Higgins & Riley (1970: 235-238) and Higgins (1975: 109-112)
for Europe under the genera Nordmannia and Strymonidia, together
with numerous related species in the central and eastern Palearctic.
The usage of Higgins & Riley, and Higgins, is representative of that
current in the Palearctic literature. Study indicates, however, that neither
of these generic names is tenable, and that assignment of the species
(i.e., which species are congeneric with which) must be modified.

Malicky (1969) has shown that the long unused generic name Fixsenia
Tutt must be revived for the widespread species pruni Linnaeus in addi-
tion to the (Asiatic) type species, herzi Fixsen. Some time ago I pointed
out (Clench 1961: 212) that pruni was congeneric with certain North
American hairstreaks to which I gave the new generic name Euristrymon.
In consequence of Malicky’s discovery, Euristrymon must therefore fall
to Fixsenia.

Malicky also concluded that other Old World hairstreaks (he specif-
ically cites spini, ilicis, w-album, and acaciae) are congeneric among
themselves, and collectively differ from Fixsenia in the presence of a
serrated ventral keel at the distal end of the penis in the male genitalia,
as well as in certain larval and pupal characters. Malicky assigned all
these species to the genus Strymonidia Tutt, a name which has long
been in general use for many of them. The name Strymonidia, however,
has as its generic type another little known species from Asia, thalia
Leech, which Malicky was not able to study. I have seen no thalia
either, but from published illustrations it seemed quite similar to herzi,
the type of Fixsenia, and I was curious about its genitalic structure.
In correspondence with Dr. Malicky I asked him about it. He was
able to borrow a specimen and sent me a drawing of the genitalic

278 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

preparation he made. This showed clearly that thalia is indeed con-
generic with herzi. Therefore the generic name Strymonidia is also a
synonym of Fixsenia and cannot be applied to the species Malicky cites.

The species Malicky included in Strymonidia, and additional Old
World species as well, are themselves congeneric with a large number
of Nearctic species now united in the genus Satyrium Scudder. This
name, Satyrium, has priority over any other and must therefore be
applied to these palearctic species.

The discovery that the numerous species of Satyrium in both Old
and New Worlds are all congeneric has created a large generic synonymy,
particularly involving the Palearctic species. The apparent reason is that
despite their close structural similarity they have diversified to a con-
siderable degree in appearance. A number of species, among them
the type of the genus, combine the un-hairstreak attributes of incon-
spicuous or absent scent pad, absence of tails and tornal lobe, and
essentially spotted patterns. They much resemble blues, and many were
originally so described. Bethune-Baker (1892), in fact, made a special
point of demonstrating that several of them were actually hairstreaks
instead of blues.

The external structural diversity of Satyrium and Fixsenia, in male
scent pad, in wing shape, in tails, and in pattern, finds a remarkable
parallel in the New World hairstreak genus Strymon Hiibner 1818 (i.e.,
as delimited in Clench 1961: 215ff). In Strymon an even more striking
genitalic homogeneity is associated with similar variation in external
structures. Apparently in both these genera extensive adaptive radiation
has occurred, although the environmental significance of most of the
affected external traits remains largely obscure.

The male genitalia of the European species of both these genera are
well figured by Higgins (1975).

The formal synonymy and nomenclatorial data for both Fixsenia and

Satyrium, together with their characters and known included species,
are as follows:

Fixsenia Tutt

Fixsenia Tutt 1907, Nat. Hist. British Butts. 2: 142, type species by original designa-
tion, Thecla herzi Fixsen 1887; Hemming 1967: 193; Malicky 1969: 38, 61.

= Leechia Tutt 1907, l.c., type species by original designation, Thecla thalia Leech
1893; Hemming 1967: 249. Junior homonym of Leechia South 1901. See
Strymonidia.

- Strymonidia Tutt 1908, op. cit.: 483, replacement name for Leechia Tutt 1907,

q.v., with the same type species; Hemming 1967: 419. New subjective synonym.

= Euristrymon Clench 1961; 212, type species by original designation, Papilio favonius
J. &. Smith 1797; Cowan 1970: 10; dos Passos 1970, J. Lepid. Soc. 24: 33. New
subjective synonym.

= Thecla, Nordmannia, etc., of authors, in part.

VoLUME 32, NuMBER 4 279

The generic characters of Fixsenia are: hindwings usually tailed, tornal lobe
usually present, if slight, but no tornal cleft. Male genitalia: uncus lobes low and
transverse, the lateral border short; vinculum with dorsal part wide, but without
posterior shoulder process, abruptly narrowing to the strap-like ventral part; anterior
border of vinculum without corematal processes; saccus present but short, rarely if
ever longer than its width at middle; valvae contiguous to more or less their middle,
then divergent, the mesial edges not dentate, and with no terminal spine; penis
apically upcurved and flared, wih two terminal cornuti of about equal diameter, but
with no terminal ventral keel.

In the Palearctic the last character, the absence of a ventral penial
keel, is sufficient to separate Fixsenia from Satyrium, the only other
genus with which it may be confused; but the remaining characters
are necessary to discriminate it from other New World Strymonine
genera. The known members are:

Palearctic species: herzi Fixsen 1887; thalia Leech 1893; pruni Linnaeus 1758.
Nearctic species: favonius J. E. Smith 1797; ontario Edwards 1868; polingi Barnes
& Benjamin 1926.

Satyrium Scudder

= Argus Gerhard 1850, Versuch. Mon. europ. Schmett. (1):4, type species by mono-
typy, Lycaena ledereri Boisduval 1848; Hemming 1967: 56. Junior homonym of
Argus Bohadsch 1761.

Satyrium Scudder 1876, Bull. Buffalo Soc. Nat. Sci. 3: 106, type species by original
designation, Lycaena fuliginosa Edwards 1861; Comstock & Huntington 1958,
J. New York Ent. Soc. 66: 116; Ziegler 1960, J. Lepid. Soc. 14: 20; Hemming
1967: 403; dos Passos 1970, op. cit.: 28.

= Callipsyche Scudder 1876, l.c., type species by original designation, Thecla behrii
Edwards 1870; Comstock & Huntington 1958, op. cit.: 105; Hemming 1967: 91.

= Neolycaena de Niceville 1890, Butts. India, Burmah and Ceylon 3: 15, 64, type
species by original designation, Lycaena sinensis Alphéraky 1881; Hemming
1967: 308.

= Edwardsia Tutt 1907, l.c., type species by original designation, Papilio w-album
Knoch 1782; Hemming 1967: 156. Junior homonym of Edwardsia Costa 1838.
See Chattendenia.

= Felderia Tutt 1907, l.c., type species by original designation, Thecla w-album
Knoch 1782; Hemming 1967: 156. Junior homonym of Edwardsia Costa 1838.
See Chattendenia.

= Felderia Tutt 1907, l.c., type species by original designation, Thecla w-album
Knoch var. eximia Fixsen 1887; Hemming 1967: 193. Junior homonym of Fel-
deria Walsingham 1887. See Thecliolia.

= Klugia Tutt 1907, l.c., type species by original designation, Papilio spini [Denis &
Schiffermiiller] 1775; Hemming 1967: 242. Junior homonym of Klugia Robineau-
Desvoidy 1863. See Tuttiola.

= Kollaria Tutt 1907, l.c., type species by original designation, Thecla sassanides
Kollar [1849]; Hemming 1967: 242. Junior homonym of Kollaria Pictet 1841.
See Superflua.

= Erschoffia Tutt 1907, l.c., type species by original designation, Thecla lunulata
Erschoff 1874; Hemming 1967: 169. Junior homonym of Erschoffia Swinhoe
1900. See Pseudothecla.

= Bakeria Tutt 1907, l.c., type species by original designation, Lycaena ledereri Bois-
duval 1848; Hemming 1967: 72. Junior homonym of Bakeria Kieffer 1905, but
never replaced.

280 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

— Nordmannia Tutt 1907, l.c., type species by original designation, Lycaena myrtale
Klug 1834; Hemming 1967: 315.

— Chattendenia Tutt 1908, op. cit.: 483; Hemming 1967: 109. Replacement name for
Edwardsia Tutt 1907, q.v., with the same type species.

= Thecliolia Strand 1910, Entomol. Rundsch. 27: 162; Hemming 1967: 439. Replace-
ment name for Felderia Tutt 1907, q.v., with the same type species.

= Tuttiola Strand 1910, l.c.; Hemming 1967: 451. Replacement name for Klugia
Tutt 1907, g.v., with the same type species.

= Superflua Strand 1910, l.c.; Hemming 1967: 420. Replacement name for Kollaria
Tutt 1907, qg.v., with the same type species.

= Pseudothecla Strand 1910, l.c.; Hemming 1967: 386. Replacement name for
Erschoffia Tutt 1907, q.v., with the same type species.

= Thecliola Waterhouse 1912, Index Zool. 2: 299; Hemming 1967: 439. Incorrect
Subsequent Spelling of Thecliolia, q.v.

= Necovatia Verity 1951, Rev. franc. Lépid., Suppl.: 183, type species by original
designation, Papilio acaciae Fabricius 1787 [proposed as a subgenus of Stry-
monidia Tutt 1908, vide supra]. Note: This generic name was overlooked by
Hemming (1967), Cowan (1968, 1970), and the Zoological Record. I thank
Lt. Col. J. N. Eliot for calling it to my attention.

= Thecla, Strymon, Strymonidia, etc., of authors (in part).

I have examined the male genitalia of the following species and find them all
congeneric:

fuliginosum, type species of Satyrium

behrii, type species of Callipsyche

w-album, type species of Edwardsia, Chattendenia
spini, type species of Klugia, Tuttiola

sassanides, type species of Kollaria, Superflua
lunulatum, type species of Erschoffia, Pseudothecla
ledereri, type species of Bakeria

myrtale, type species of Nordmannia

acaciae, type species of Necovatia

I have not seen the species sinensis (type species of Neolycaena),
but pretiosum Staudinger is extremely closely related, perhaps conspe-
cific, and its genitalia, along with those of some other Satyrium, were
figured by Bethune Baker (1892). His figures show that pretiosum,
and hence most likely sinensis, is a Satyrium. I have not seen eximium
Fixsen (type species of Felderia, Thecliolia), but from illustrations it
seems to be a close relative of w-album and I conclude, therefore, that
it is probably congeneric.

The generic characters of Satyrium are: male genitalia: distal end of penis with
ventral serrated keel; penis with two terminal cornuti, one of which is usually dentate;
distally divergent valvae; tips of valvae without a mesial hair-like fringe.

In the Palearctic the presence of a serrated penial keel will discrim-
inate it from Fixsenia, or any other Strymonine. In the New World,
however, several other genera share this keel, from which the additional
characters will separate it. The known species belonging to Satyrium
are as follows. I include only those whose genitalia I have examined
either directly or in published illustrations, except for the species fol-

VoLUME 32, NuMBER 4 281

lowed by “(?),” which are included provisionally, on the basis of ex-
ternal facies alone.

Palearctic species: eximium Fixsen 1887 (?); w-album Knoch 1782;
spini [Denis & Schiffermiiller] 1775; latior Fixsen 1887; sassanides Kollar
1849: lunulatum Erschoff 1874; pretiosum Staudinger 1886; sinensis
Alpheéraky 1881 (?); ledereri Boisduval 1848; myrtale Klug 1834; teng-
stroemii Erschoff 1874; ilicis Esper 1779; acaciae Fabricius 1787.

Nearctic species: fuliginosum Edwards 1861; behrii Edwards 1870;
auretorum Boisduval 1852; saepium Boisduval 1852; tetra Edwards 1870
(= adenostomatis Hy. Edwards 1877); liparops Le Conte 1833; kingi
Klots & Clench 1952; calanus Hiibner 1809; caryaevorum McDunnough
1942; edwardsii Scudder 1870; sylvinum Boisduval 1852; californicum
Edwards 1862; acadicum Edwards 1862.

It is worth noting that at one time it was believed (e.g., Riley 1958:
285) that myrtale was congeneric with Erora Scudder 1872. The latter
genus, however, is representative of a wholly New World, primarily
tropical, group with no palearctic members at all. This group differs
considerably from Satyrium, not only in the absence of a serrated penial
keel, but also in the high lateral margins of the uncus lobes, the single,
always interior, cornutus, and in addition a terminal penial tooth that
is either external and part of the shaft or else internal, on the vesica,
and eversible. The vesica, moreover, is usually scobinate.

LITERATURE CITED

BETHUNE-BAKER, G. T. 1892. Notes on Lycaena (recte Thecla) rhymnus, teng-
stroemii, and pretiosa. Trans. Entomol. Soc. London 1892: 27-31, ill.

Ciencu, H. K. 1961. Tribe Theclini, pp. 177-220, in P. R. and A. H. Ehrlich,
How to know the butterflies. W.C. Brown, Dubuque, Iowa.

Cowan, C. F. 1968. Annotationes Rhopalocerologicae. Author, Berkhamsted,
England. 20 pp.

1970. Annotationes Rhopalocerologicae 1970. Berkhamsted, England:
author; 70 pp.

Hemminc, F. 1967. The generic names of the butterflies and their type-species
(Lepidoptera: Rhopalocera). Bull. Br. Mus. Nat. Hist. (Entomol.), Suppl. 9:
509 pp.

Hiccrns, L. G. 1975. The classification of European butterflies. Collins, London.
320 pp., ill.

& N. D. Rmey. 1970. A field guide to the butterflies of Britain and Europe.
Collins, London. 380 pp., ill.

Mauicxy, H. 1969. Uebersicht ueber praimaginalstadien, bionomie und O6kologie
der mitteleuropaischen Lycaenidae (Lepidoptera). Mitt. Entomol. Ges. Basel
(N.F.) 19: 25-91, ill.

Ritty, N. D. 1958. The genera of holarctic Theclinae: A tentative revision. Proc.
Tenth Int. Congress Entomol. 1: 281-288.

Journal of the Lepidopterists’ Society
32(4), 1978, 282-288

OVER-WINTERING BEHAVIOR IN EUPHYDRYAS PHAETON
(NYMPHALIDAE)

M. DEANE BOWERS
Department of Zoology, University of Massachusetts, Amherst, Massachusetts 01003

ABSTRACT. The behavior of the pre-diapause larvae of Euphydryas phaeton
(Nymphalidae) associated with over-wintering is described. Observations are based
on studies of five wild populations in central Massachusetts over a three-year period,
as well as on larvae reared in the laboratory. The functional significance of this
larval behavior is discussed.

Most Lepidoptera over-winter as either an egg or pupa; some nym-
phalid butterflies (e.g. the Mourning Cloak, Nymphalis antiopa L., and
Compton’s Tortoiseshell, N. vau-album Boisduval and Leconte), and
certain groups of moths [e.g. the Lithophanini (Schweitzer, 1977) |
overwinter as adults. In only a relatively few cases do Lepidoptera
spend the winter in the larval stage. The Viceroy, Limenitis archippus
Cramer (Nymphalidae) for example, is multivoltine and may enter a
facultative diapause in the third instar during late summer and fall
(Hong and Platt, 1975). Unusually, however, the Baltimore Checker-
spot, Euphydryas phaeton Drury (Nymphalidae), and other members
of the genus Euphydryas, are univoltine and exhibit an obligatory dia-
pause in the fourth instar.

Although some descriptive work has been done on the development
of the early, pre-diapause instars of E. phaeton, little has been reported
of the actual behavior of the larvae during over-wintering. Observa-
tions on several wild colonies of E. phaeton and on larvae reared. in
the laboratory have revealed some interesting behavioral aspects of the
over-wintering process in this species.

METHODS

Observations were made on five colonies of E. phaeton in Hampshire,
Franklin, and Hamden Counties in central Massachusetts from 1974—_
1977. The site of each colony was wet and marshy, typical habitat for
E. phaeton and its primary foodplant, Turtlehead (Chelone glabra L..,
Scrophulariaceae). The five sites differed in elevation, area, amount
of Turtlehead present, and size of the E. phaeton population; but these
differences were not factors in the present study.

At each of the colonies, I determined the absolute numbers of egg
masses, and (later) larval webs. At the three largest colonies, heights
of the pre-hibernation webs above the ground were measured. All

VoLUME 32, NuMBER 4 283

colonies were usually observed at least once a week. In addition I
reared larvae from eggs in the laboratory at 22-25°C under continuous
light. Developmental times for each instar, dates of larvae entering
the pre-hibernation web and ceasing feeding, and time to the third
molt were recorded for these larvae.

OBSERVATIONS

Euphydryas phaeton females usually oviposit two to three large masses
of from 100-600 eggs each (Scudder, 1889; Bowers, pers. obs.) from late
June through July in Massachusetts. The eggs hatch in about 20 days.
When the larvae hatch they may construct a small web on the leaf next to
the egg shell but move to the growing tip of the plant within 24 hours
and begin web construction and feeding. This feeding web is extended
down the length of a stalk of Turtlehead as the larvae devour the leaves.
The larvae are gregarious and develop to the end of the third instar in
approximately three weeks.

About mid-August in central Massachusetts, the larvae stop feeding,
and thicken and compact a section of the web (this date may vary
widely, depending on the hatch date of a particular cohort of larvae,
elevation, and current climatic conditions). Upon entering this “pre-
hibernation web” larvae become quiescent and molt to the fourth instar
in about five days (from 3-7 days in the laboratory) (Edwards, 1875;
Scudder, 1889; Bowers, pers. obs.). Edwards (1875) reported that larvae
enter this web about 15 July in West Virginia, but at higher elevations in
Massachusetts this web entrance can occur as late as the second week
in September.

Previous authors (e.g. Scudder, 1889; Clark, 1927) have referred to
this strengthened and compacted web as the “hibernating web.” How-
ever, larvae do not spend the winter in this web at all, but leave it
and move into the litter; this web is thus better referred to as the
“pre-hibernation web,” which is the term used throughout this paper.
This designation also serves to differentiate it from the feeding web
which encloses plant parts on which the larvae are feeding.

The pre-hibernation web may be constructed at the base of the
Turtlehead stalk on which the larvae have been feeding, but is some-
times found a short distance away. This web, as well as the feeding
web constructed throughout the development of the pre-hibernation
larvae, includes the stalk and leaves of Turtlehead as well as adjacent
plants such as ferns, grasses and herbs (Figs. 1-3). The pre-hibernation
webs are always constructed above ground level; most are found at a
height of more than 50 cm. The mean height for 43 pre-hibernation

284 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

VoLUME 32, NUMBER 4 285

webs measured in the summer of 1976 was 73 cm (range: 30-127 cm;
standard deviation: + 25.50). I have never observed a pre-hibernation
web directly on the ground.

Once the larvae enter the pre-hibernation web they do not feed at
all. There may be some movement and occasionally a larva or two
will leave and wander over the web surface. If the larvae are disturbed
by movement of the web, they will move around a bit, but are usually
inactive. An exception to this occurs when the web is damaged by
storms or predators. In order to observe larval reactions to such damage,
I broke open several webs. When the web is broken, the larvae begin
moving around almost immediately and one or two larvae begin to
crawl out of the web and over the surface. Other larvae soon follow.
Within a day or two the web has been repaired and the larvae return
to their inactive state. Thus, although the larvae are quiescent and
not feeding, they can still react to stimuli such as disruption of the
web.

The larvae do not spend the winter inside this web as has been
suggested by previous authors (e.g. Edwards, 1875; Scudder, 1889;
Clark, 1927). Rather, about the end of October, the larvae move out
of the web en masse to the base of a plant on which the pre-hibernation
web was constructed. Here they form a large contiguous aggregation
among the dead grass and litter on the ground. This movement occurred
during the weeks 27 October-4 November 1975, 22 October-29 October
1976, and 20 October-27 October 1977. While in this aggregation, in-
dividual larvae are active on warm days, moving from the primary mass
as far as 25 cm. On cool, cloudy days or early in the morning, the
larvae are usually found in a tight group, but individuals will quickly
move around and out of the mass when disturbed. After approximately
a week (from two days to two weeks; different for each aggregation
of larvae), groups of about 10 to 100 individuals from this large mass
move distances of 5-100 cm away and roll up leaves and bits of debris
and fasten them with silk. It is here that they spend the winter. From
1974 to 1977 over 100 pre-hibernation webs were examined and in all
webs the larvae moved into the litter. All the larvae remain near the
pre-hibernation web and thus close to the Turtlehead stalk on which

<

Figs. 1-3. Development of the feeding web of E. phaeton on C. glabra: 1,
beginning web formation at top of plant; 2, further development of web with more
leaves enclosed; 3, extension of web to encompass several stalks of C. glabra and
adjacent plants.

286 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

they originally fed. Because Turtlehead is perennial this behavior en-
sures an adequate food supply for larvae emerging the following spring.

On warm, sunny days during the winter, larvae may become active,
and groups may shift position by a few cm, rolling up new leaves in
which to spend the remainder of the winter.

When the larvae are dug up from under the snow in January and
brought to temperatures of 20-25°C, they quickly become active and
crawl around the container in which they are confined. Usually they
will not feed at this time. Clark (1927) brought hibernating larvae
indoors in February and tried feeding them forced shoots of Lonicera
japonica Thunb. (Caprifoliaceae ), an alternate foodplant for this species
(Clark, 1927; Scudder, 1889). After wandering around for a week,
they began to feed, but all eventually died before reaching pupation.
If larvae are brought indoors in this manner, allowed to be active and
then returned to outside conditions, they resume their resting state
with no apparent ill effects, emerging from diapause in the normal
manner in the spring.

I attempted to break diapause in E. phaeton by subjecting several
egg masses and their emerging larvae to 24 hours of light throughout
development and hibernation. Although little is known about breaking
diapause in univoltine insects, Beck (1968) suggests this method. These
lab-reared larvae formed a pre-hibernation web in the same way as
larvae in the field, yet were occasionally active. The larvae were given
fresh food when plants seemed wilted or were consumed, and _ they
fed occasionally through the end of October, with approximately 40%
mortality. Between 25 October to 1 November there was a fourth molt
(which does not occur in natural populations) that was synchronized
within any single group of larvae but not among the groups. After
this time, most of the larvae were inactive, and by mid-January all the
larvae had died. Although attempts to break diapause in other insects
with an obligatory diapause have been successful (e.g. Beck, 1968),
this attempt was not. None of the larvae survived and none exceeded
the size attained by hibernating larvae in the field. Thus diapause in
this univoltine insect is truly obligatory and probably under genetic
rather than photoperiodic control; the latter is probably the case in
most multivoltine species (Beck, 1968; Hong and Platt, 1975).

DISCUSSION

Karly authors (e.g. Scudder (1889) and Edwards (1884)) believed
that the larvae of E. phaeton spent the winter in the pre-hibernation
web (which they called the “hibernation web”). Observation of these

VoLUME 32, NUMBER 4 287

webs throughout the fall in Massachusetts shows that it would not be
feasible for larvae to remain in the web: although the pre-hibernation
web has many more layers of silk and is smaller and more compact
than the feeding web, wind and rain begin to damage it by the end
of October. Before winter has advanced very far there is little left of
the web, and the larvae cannot repair the web during the winter.

One of the harshest winter microclimates is just above the snow level
where most of the pre-hibernation webs are found. Air temperature
fluctuations here are much more dramatic than those at the soil surface
(MacKinney, 1929; Mail, 1932) where larvae aggregate. Vegetation
and litter reduce these temperature fluctuations, and snow provides
further insulation (MacKinney, 1929; Mail, 1932). In Massachusetts
there is snow cover for most of the winter and thus the larvae are
quite well protected. Wind is also an important agent above the snow
or ground level, but would have little impact on the larvae under
snow or litter cover. Thus, regardless of snow cover, by moving out
of the pre-hibernation web into the litter, larvae escape extreme tem-
perature fluctuations and the desiccating effects of wind characteristic
of winter conditions.

The question, then, is why should larvae construct a pre-hibernation
web at all; why not move into the litter in August when feeding stops?
Perhaps the groups of hibernating larvae would be easy prey for ground
predators such as spiders and beetles which are abundant at the end
of the summer. By the end of October, however, most of these pred-
ators are absent or inactive and the larvae could safely move into the
litter.

In summary, pre-hibernation larvae of E. phaeton exhibit three major
behavioral sequences: first, construction of the feeding web during the
first three instars; second, abandonment of this web and construction
of a small, compact pre-hibernation web in which the larvae remain
quiescent; third, departure from this web and movement of smaller
groups into the litter where they form an over-wintering site by rolling
up leaves and bits of debris and fastening them with silk. The last two
sequences require appreciable expenditures of energy (for movement
and silk-making) while no food is being eaten. This expenditure sug-
gests that these behaviors are necessary to ensure larval survival over
the fall and winter.

ACKNOWLEDGMENTS

I am very grateful to Dr. Theodore D. Sargent for his encouragement
and editorial comments. This research was supported in part by a
Sigma Xi Grant-in-Aid of Research.

288 JoURNAL OF THE LEPIDOPTERISTS SOCIETY

LITERATURE CITED

Beck, S. D. 1968. Insect photoperiodism. Academic Press, New York. 288 pp.

Criark, A. H. 1927. Notes on the melitaeid butterfly Euphydryas phaeton (Drury )
with descriptions of a new subspecies and a new variety. Proc. U.S. Nat. Mus.
Wash. 71, article 11, #2683. pp. 1-22.

Criark, S. H. & A. P. Puarr. 1969. Influence of photoperiod on development and
larval diapause in the Viceroy butterfly, Limenitis archippus. J. Insect Physiol.
15: 1951-1957.

Epwarps, W. H. 1875. Notes on butterflies. Canad. Entomol. 7: 150-151.

1884. Description of the preparatory stages of Melitaea chalcedon, Bois.,
with some notes on larvae of M. phaeton. Papilio 4: 63-70.

Hone, J. W. & A. P. Puarr. 1975. Critical photoperiod and day-length threshold
differences between northern and southern populations of the butterfly, Limenitis
archippus. J. Insect Physiol. 21: 1159-1165.

MackKinney, A. L. 1929. Effects of forest litter on soil temperature and soil freez-
ing in autumn and winter. Ecology 10: 312-321.

Mart, G. A. 1932. Winter temperature gradients as a factor in insect survival. J.
Econ. Entomol. 25: 1049-1053.

ScuppER, S. 1889. The Butterflies of the Eastern United States and Canada. W.
H. Wheeler, Cambridge, Mass.

ScHweitzER, D. F. 1977. Life history strategies of the Lithophanini (Lepidoptera:
Noctuidae, Cuculliinae), the winter moths. Ph.D. dissertation. Univ. of Mass.,
Amherst, Mass. 304 pp.

Journal of the Lepidopterists’ Society
32(4), 1978, 289-303

INTER-SPECIFIC HYBRIDIZATION INVOLVING LIMENITIS
ARCHIPPUS AND ITS CONGENERIC SPECIES
(NYMPHALIDAE)

AUSTIN P. PLATT

Department of Biological Sciences, University of Maryland Baltimore County,
5401 Wilkens Avenue, Catonsville, Maryland 21228

GrorGE W. RAWSON
10405 Amherst Avenue, Silver Spring, Maryland 20902

GEORGE BALOGH
3607 No. 98th Street, Milwaukee, Wisconsin 53222

ABSTRACT. The occurrence of 43 natural hybrids involving Limenitis archippus
and its congeneric species (L. arthemis-astyanax, L. lorquini, and L. weidemeyezrii )
is reviewed. Nine of these hybrid records are reported for the first time. Data
based on laboratory crosses are given in order to document the purported wild hybrid
specimens. Reasons underlying the observed natural hybridization are suggested and
their evolutionary implications are discussed.

Species of the North American genus Limenitis readily undergo inter-
specific hybridization both in nature and in the laboratory (Edwards,
1882: Scudder, 1889; Field, 1904, 1914; Newcomb, 1907; Gunder, 1934;
Remington, 1958, 1968; Platt, 1975). The occurrence of 22 natural
hybrids and the laboratory documentation of them in crosses involving
either 1) L. arthemis arthemis Drury or 2) L. arthemis astyanax Fabri-
cius X L. archippus Cramer have been reviewed and discussed by
Monroe (1953); Grey (1968); Shapiro and Biggs (1968); Platt and
Greenfield (1971), and Greenfield and Platt (1974). Since then, John-
son (1974) and Arbogast (1976) have reported two other wild-collected
L. arthemis astyanax X L. archippus hybrid specimens. Likewise, the
natural occurrence of four L. lorquini Boisduval x L. archippus hybrids
(Gage, 1970; Perkins and Gage, 1970) and five L. weidemeyerii Ed-
wards X L. archippus hybrids (Cross, 1936, 1937; Simpson and Pettus,
1976) have also been recorded. Lab-bred equivalents of these wild
hybrids are shown in Fig. 1.

In this paper we shall review these past records and will report
records of nine other naturally occurring Limenitis hybrids involving
L. archippus, a species broadly sympatric with other members of the
genus. We will also present new information obtained from laboratory

bo
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JouRNAL OF THE LEPIDOPTERISTS SOCIETY

DORSAL VENTRAL

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Fig. 1. Representative lab-bred F, male hybrid specimens. 1) Form “arthechip-
pus, brood 893, No. 17, May 7, 1977; 2) form “weidechippus,” brood 576, No. 7,
Sept. 4, 1971; 3) lorquini 9 x archippus 6 (unnamed hybrid), brood 987, No. 1,
Sept. 8, 1978. These, and all other lab-bred specimens, were reared on either Salix
babylonica L. or Prunus serotina Ehrh.

VoLUME 32, NuMBER 4 291

UNITED STATES

$000", 10000" contours

“EH. Frowschnar ond BLE. Woodion. Jr

Fig. 2. Distribution map of 43 known wild, inter-specific F; hybrids involving
cross-breeding with L. archippus collected from “prior to 1872” through 1976. All
records for which the sex is known are males. These hybrids are widely distributed
geographically. Touching symbols represent two (or more) specimens from the same
locality. Most hybrids have been collected late in the season ( August-November ).
Complete data for these hybrids are given in Tables 1, 1A, and 2. (Map reproduced
with the permission of the Missouri Botanical Garden, St. Louis, Mo. ).

crosses recently made by Platt, followed by a brief discussion of the
relationships between L. archippus and its close relatives.

Tables 1, [A and 2 summarize the collection data for all 43 records of
wild hybrids involving L. archippus and its congeneric species. The
geographic distributions of the various hybrid forms have been plotted
in Fig. 2. The new hybrid locality records given in Table 1 are those
from Maine (hyb. form “arthechippus” Scudder), Wisconsin, Tlinois,
Michigan, New Jersey, Virginia, and Florida (hyb. form “rubidus”
Strecker). The specimens from Idaho, Illinois, Michigan, New Jersey,
Virginia, and Florida probably represent state records for these hybrids.

The Florida record represents the first report of a natural hybrid
between L. a. astyanax Fabricius and the sub-species L. archippus
floridensis Strecker. The live specimen was observed circling a shrub
willow along the edge of Mud Lake by G.W.R. while he was lunching

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Fig. 3. 1) Wild-caught and 2) lab-bred male specimens of L. arthemis astyanax
x L. archippus floridensis F1 hybrids (form “rubidus”). Data for the lab-bred speci-
men are brood 986, No. 1, Sept. 19, 1977 (Data for wild-caught specimen are given
in Table 1).

with members of the Florida Audubon Society in the northeastern
portion of the Ocala National Forest. He recognized the specimen
as an “off-color” Limenitis and investigated it further. The insect then
settled on the willow shrub. Since he had no net, George made a
“desperate strike” at it with his cap, knocking the butterfly to the ground
and collecting it. The specimen is illustrated in Fig. 3, along with a
similar, single specimen recently reared by A.P.P. in the laboratory
from a hand-paired cross between a Maryland astyanax ? X a floridensis
é (from a stock obtained just east of the Everglades [near Home-

VoLUME 32, NuMBER 4 297

DORSAL VENTRAL

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Fig. 4. Small, weak, and faded lab-bred F, female hybrid specimens, dorsal and
ventral views. 1) Form “weidechippus,” brood 576, No. 1, Aug. 28, 1971; 2) form
“rubidus,’ brood 648, No. 4, July 26, 1973.

stead] in southern Florida). Astyanax evidently is relatively uncommon
in central Florida. Near his home at New Smyrna Beach (Volusia Co.),
G.W.R. has seen only two specimens of astyanax during the past 20
years. Also, F. Rutkowski (pers. comm.) recently collected a ¢ astyanax
1.5 miles north of Shamrock (Dixie Co.) in Florida, and he mentioned
other records from as far south as Dade Co. (Kimball, 1965). Possibly
such scarcity of one (or both) species accounts in part for such inter-
specific hybridization (Simpson and Pettus, loc. cit.).

So far as is known, all of the wild hybrids thus far collected have
been males, although five broods containing small, faded, weak (and
often malformed) F, females (Fig. 4) were reared at UMBC in June
and July, 1973 by crossing inter-specific strains having different geo-
graphic origins (Maryland L. astyanax 22 X Vermont L. archippus

DORSAL

JouRNAL OF THE LEPIDOPTERISTS SOCIETY

VENTRAL

VoLUME 32, NUMBER 4 299

é 4). Among these broods, there were a total of 39 (27%) F, females
among 143 hybrid “rubidus” progeny. Two of the larger broods, in-
volving sibling female astyanax and the same male archippus parent,
yielded 1:1 sex ratios.

Earlier crosses reported by Platt (1975) showed that such inter-
specific crosses, in which strains from the same (or closely adjacent)
northeastern geographic origins were crossed, gave rise to complete
adult heterogametic (female) inviability. However, robust females of
the “arthechippus” and “rubidus” hybrid phenotypes (Fig. 5) also have
been obtained by backcrossing F, hybrid males to females of the three
parental forms (arthemis, astyanax, and archippus, respectively). Such
backcrosses often have low viability, but sometimes yield relatively
large numbers of progeny (Platt, loc. cit.). Although fertile crosses
have been obtained in all possible reciprocal combinations, crosses using
L. arthemis, astyanax, lorquini, or weidemeyerii 22 X L. archippus 3 4
have been the easiest to effect in the laboratory using hand-pairing.
The fact that such a pairing also has been seen in the wild (Klots,
1959; Table 1A) suggests that these inter-specific hybrid crosses may
occur most often in this direction in nature, as well.

Table 1 indicates that eight of the nine previously unreported arthemis-
astyanax X archippus hybrids, like most of those reported earlier, were
collected during the late summer and fall months (August-November).
Only one (from New Jersey) was collected in June, a time suggesting
that it most likely arose from an over-wintering larva. Thus, these new
records, as well as the previous ones, support the contention that the
ecological and behavioral barriers normally preventing inter-specific
hybridization in Limenitis tend to break down later in the season (Green-
field and Platt, loc. cit.). This break down may well be correlated with
the onset of facultative larval diapause in Limenitis which occurs during
the third instar. Thus, it seems as if those individuals most often
selecting mates of the wrong species are the very ones which seem to
be “genetically mal-adapted” to their environment (that is, they are the
ones which are not diapausing at that time of year when they are

<&

Fig. 5. Representative robust, lab-bred hybrid-type backcross females. 1) L.
arthemis 2 xX F, hybrid “arthechippus” 8 (“arthechippus-like” morph), brood 63c,
No. 4, July 30, 1968; 2) L. archippus 2 xX F; hybrid “arthechippus” ¢ (“arthe-
chippus-like” morph), brood 915, No. 45, June 13, 1977; 3) L. arthemis 2 xX F;
hybrid “rubidus’ 8 (“proserpina-like” morph), brood 95B, No. 4, Dec. 26, 1968;
4) L. archippus 2 & F; hybrid “rubidus” ¢ (“rubidus-like” morph), brood 757, No.
10, Sept. 15, 1975. Such backcross females also occur in parent-type morphs (see
Pia LO):

JoURNAL OF THE LEPIDOPTERISTS SOCIETY

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supposed to be doing so). Such hybridization, perhaps, represents a
“last chance” effort to reproduce.

The similar rare natural hybrids reported between both of the western
banded Limenitis (L. lorquini and L. weidemeyerii) and L. archippus
are listed in Table 2. The eight known dates of siting or capture sug-
gest once again that either 1) the hybrids emerged early enough in
the year (June and July dates) so that it is a relative certainty that
the larvae from which they arose over-wintered in hibernacula, or 2)
the specimens were collected in the late summer or fall months. Gage’s
(pers. comm.) collection dates for the small “hybrid swarm” near Rich-
land, Washington (Table 2) suggest that the four lorquini x archippus
hybrids, in fact, represent progeny from at least three different matings.
The same may be said for the four “weidechippus” records from Colo-
rado, as well. All of these wild western hybrids are males, and they
closely resemble hybrid from “arthechippus” in possessing a_ partial
postmedial white band dorsally (Fig. 1). Two crosses between Colo-
rado weidemeyerii 22 X Massachusetts archippus 6 ¢ have been made
by laboratory hand-pairings to date, yielding 22 ¢¢ and nine 2°
(seven of the latter being malformed). All of these F,’s, although
showing some phenotypic variability, are referrable to hyb. form “weide-
chippus’ (Platt, unpub. data). During the past summer two crosses
between Oregon L. lorquini °° X Maryland L. archippus ¢é were
carried out. All 16 F, progeny were males. An additional cross involving
an F, hybrid, arthemis-lorquini 2 (Massachusetts = Oregon stocks,
respectively) x Maryland archippus ¢, yielded 21 male hybrid-like
progeny.

Thus, L. archippus, which is broadly sympatric with its congeners,
will occasionally hybridize with all of the other allopatric species of
Limenitis in nature. However, such crosses evidently are rare, leading
to the supposition that morphological, behavioral, visual, and possibly
pheromonal cues, as well as habitat isolating mechanisms, normally
operate to prevent such inter-specific hybridization. These barriers
against gene exchange between the viceroy and its close relatives some-
times tend to break down, usually when one or both species are rare,
and often toward the end of the breeding season, at times when the
majority of developing Limenitis larvae are entering diapause.

Laboratory data show that inter-specific strains having different geo-
graphic origins may be genetically more compatible than similar strains
from the same locality, as judged by either the presence or absence of
adult females in the F, generation. Thus, genetic incompatibility be-
tween the viceroy and its congeneric species is viewed as being of local

302 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

origin, suggesting that archippus may have arisen from a banded an-
cestral species by the process of sympatric speciation. (The senior
author would appreciate hearing from members of the Society who
may have knowledge of other records of wild Limenitis hybrids. )

ACKNOWLEDGMENTS

We are grateful to those individuals who provided us with informa-
tion regarding the collection of hybrid specimens as noted in Table 1.
We thank Dr. A. Maizels for sending the floridensis stock and Mr. P.
J. Kean for making the inter-specific hand-pairing involving this strain.
Mr. S. J. Harrison and Mr. T. Williams have assisted with the insect
rearing and preservation. We thank Dr. W. D. Field, Dr. D. C. Ferguson
of the U.S.N.M., and Dr. K. Bagdonas of the University of Wyoming
for the information relating to hyb. form “weidechippus’. Mr. E. Gage
kindly provided data and color photos of the lorquini xX archippus
hybrids.

LITERATURE CITED

Arsocast, R. T. 1976. Capture of a hybrid Limenitis arthemis astyanax x L.
archippus (Nymphalidae) in southern Georgia. J. Lepid. Soc. 30: 4.

Cross, F.C. 1936. (Notitle). Hobbies 41: 112.

1937. Butterflies of Colorado. Proc. Colo. Mus. Nat. Hist. 16: 3-28.

Epwarps, W. H. 1882. Descriptions of new species of butterflies found in the
United States. Papilio 2: 45-49.

Frecp, W. L. W. 1904. Problems in the genus Basilarchia. Psyche 11: 1-6.

. 1914. Hybrid butterflies of the genus Basilarchia. Psyche 21: 115-117.

Gace, E. V. 1970. A record of a naturally occurring Limenitis hybrid (Nymphal-
idae). J. Lepid. Soc. 24; 270.

GREENFIELD, J. C., Jn. & A. P. PLatr. 1974. Report of the capture of an additional
hybrid between Limenitis arthemis astyanax and L. archippus (Nymphalidae).
J. Lepid. Soc. 28: 72—75.

Grey, L. P. 1968. (No title). In No. Amer. Ann. Summary, News Lepid. Soc., No.
a, p, 29)

Gunver, J. D. 1934. A check list revision of the genus Basilarchia Scud. (Lepid.:
Rhopalocera ). Canad. Entomol. 66: 39-48.

Jounson, K. 1974. An aberrant interspecific hybrid of Limenitis (Nymphalidae )
from Wisconsin. J. Lepid. Soc. 28: 163-165.

KimBati, C. P. 1965. The Lepidoptera of Florida, in Arthropods of Florida and
neighboring land areas. Fla. Dept. Agric., Gainesville, Fla. Vol. 1: 363 pp.

Kiors, A. B. 1959. A mixed mating of two species of Limenitis Fabricius (Lepidop-
tera, Nymphalidae). J. N.Y. Entomol. Soc. 67: 20.

Monroe, B. L. 1953. <A hybrid Limenitis. Lepid. News 7: 53.

Newcoms, H. H. 1907. Description of a new variety of Limenitis ursala. Psyche
14: 89-91.

Perkins, h. M. & E. V. Gacr. 1970. On the occurrence of Limenitis archippus «
L. lorquini hybrids (Nymphalidae). Journ. Res. Lepid. 9(4): 223-226.

Prarr, A. P. 1975. Monomorphic mimicry in nearctic Limenitis butterflies: experi-

mental hybridization of the L. arthemis-astyanax complex with L. archippus.
Evolution 29: 120-141.

VoLUME 32, NUMBER 4 303

Piatt, A. P. & J. C. GREENFIELD, Jr. 1971. Inter-specific hybridization between
Limenitis arthemis astyanax and L. archippus (Nymphalidae). J. Lepid. Soc.
94; 278-284.

REMINGTON, C. L. 1958. Genetics of populations of Lepidoptera. Proc. Tenth Int.
Congr. Entomol. 2: 787-805.

. 1968. Suture-zones of hybrid interaction between recently joined biotas.
Evol. Biol. 2: 321-428.

ScuppEer, S. H. 1889. The butterflies of the eastern United States and Canada,
with special reference to New England. Vol. 1. Publ. by the author, Cambridge,
Mass. pp. 250-305.

SuHapiro, A. M. & J. D. Biccs. 1968. A hybrid Limenitis from New York. J. Res.
Lepid. 7: 149-152.

Srmpson, R. G. & D. Petrus. 1976. Records of Limenitis hybrids from Colorado.
J. Res. Lepid. 15(3): 163-168.

Journal of the Lepidopterists’ Society
32(4), 1978, 303-304

TAENIDIA INTEGERRIMA, A NEW FOODPLANT RECORD FOR
PAPILIO POLYXENES (PAPILIONIDAE)

Host plants recorded for the larval stages of Papilio polyxenes Fabricius include a
wide variety of species in the family Umbelliferae. Although the dominant foodplants
in the northeastern United States are plants naturalized from Europe, e.g., Daucus
carota Linnaeus and Anethum graveolens L. (Tyler, 1975, The Swallowtail Butterflies
of North America, Naturegraph Publishers, Heraldsburg, CA), a number of endemic
species have been documented as foodplants. Tietz (1972, An Index to the Described
Life Histories, Early Stages, and Hosts of the Macrolepidoptera of the Continental
United States and Canada, A. C. Allen, Sarasota, FL) lists Cicuta bulbifera L., Cicuta
maculata L., Angelica atropurpurea L., Osmorhiza claytoni (Michx.), Osmorhiza longi-
stylis (Torr.), Oxypolis filiformis (L.), Spermolepis divaricata (L.), Ptilimnium capil-
laceaum (Michx.), and Sium suave Walt. among the native umbellifers; Cryptotaenia
canadensis (1..) has recently been reported as a foodplant as well (Scriber and Finke,
1978, J. Lepid. Soc. 32: 236-238). The majority of these species are characteristically
found in rich damp woods (Osmorhiza spp.) or wet thickets and swamps.

The native umbellifer Taenidia integerrima (L.) Drude (yellow pimpernel ), hitherto
unrecorded as a host plant for P. polyxenes but reported as a host plant of the recently
described sibling species P. joanae (Heitzman, 1973, J. Res. Lepid. 12: 1-10), is a
plant of dry, gravelly slopes and rocky hillsides (Fernald, 1950, Gray’s Manual of
Botany, 8th ed., American Book Co., NY). On June 22, 1977, one fifth instar larva
of P. polyxenes was found feeding in a patch of T. integerrima growing on a dry,
exposed slope bordering a road which cuts through Coy Glen, a forested area 3 km
west of Ithaca, Tompkins Co., New York. Two additional fifth instar caterpillars
were found in the same patch two days later. All three caterpillars were collected
and reared to pupation on T. integerrima collected from the Coy Glen site. Pupation
was virtually synchronous on June 24, 1977, indicating that the caterpillars may have
developed from eggs laid at approximately the same time, possibly by a single female.
On July 2, 1977, one adult male ichneumonid, Trogus pennator (Fabr.), a well-known
parasitoid of P. polyxenes (Heinrich, 1964, Canad. Ent. Suppl. 29: 807-853), emerged
from each of the three pupae.

304 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Scriber (1975, doctoral dissertation, Cornell University) reports that, under labora-
tory conditions, T. integerrima supports the growth of P. polyxenes as well as (or
better than) most of its commoner foodplants. Moreover, females confined over the
plant oviposit freely on it (M. Berenbaum and M. Rausher, pers. obs.). The dearth
of records for P. polyxenes on yellow pimpernel in the wild is therefore intriguing.
Habitat unsuitability might be one explanation. P. polyxenes does not normally fly
in deep woods (Tyler, 1975, op. cit.), where T. integerrima is found, and thus is not
likely to encounter the plant. In this particular instance, the plants were growing
along a road cut through thick woods approximately 0.4 km from an open field sup-
porting a luxuriant growth of Conium maculatum, a well-documented host plant for
P. polyxenes (Tietz, 1972, op. cit.). Road cuts are known to act as flyways for P.
polyxenes (Heitzman, 1973, op. cit.) and, in this case, might have directed an ovi-
positing female out of open field habitat into deep woods, where she might otherwise
not fly.

Especially interesting with respect to these findings are two other reports of P.
polyxenes on plants in heavily wooded areas. Scriber and Finke (1978, op. cit.) ob-
served an oviposition on Cryptotaenia canadensis growing in a narrow wooded strip
in an otherwise open residential area, and Rehr (1973, J. Lep. Soc. 27: 237-238)
reported the occurrence of P. polyxenes larvae on another deepwoods plant, Thaspium
barbinode ( Michx.), which was growing along a road cut on a dry bank. T. barbinode,
like Taenidia integerrima, is a purported host of the newly described P. joanae
(Tyler, 1973, op. cit.). In view of the ability of P. polyxenes to exploit both P. joanae
hostplants and habitat, and given the enormous variability in both larval and adult
coloration within the species (M. Berenbaum, W. Blau, L. Contardo and P. P. Feeny,
pers. obs.), it might well be premature to assign species status to P. joanae without
first examining the extent to which habitat has contributed to reproductive isolation
in areas where P. joanae is reported to occur.

I thank Ken Sandlan of the Cornell University Entomology Department for identi-
fying the ichneumonids (deposited along with the P. polyxenes chrysalids in Cornell
University Collection Lot 1023, Sublot 41a) and W. Blau and P. P. Feeny of the
Department of Ecology and Systematics at Cornell for confirming the identification
of the caterpillars. This work was supported in part by N.S.F. Research Grant DEB
76-20114 AOI to P. P. Feeny.

M. BerensaumM, Department of Entomology, 110 Insectary, Cornell University,
Ithaca, NY 14853

Journal of the Lepidopterists’ Scciety
32(4), 1978, 304-305

MID-VALVAL FLEXION IN THE LEFT VALVA OF ASYMMETRIC
GENITALIA OF ERYNNIS (HESPERIIDAE )

The left and right sides of Erynnis (Erynnides) Burns genitalia often differ grossly
in shape (Burns, 1964, Univ. Calif. Pub. Entomol. 34). On a hilltop at Red Rocks,
Jefferson Co., Colorado on 21 June 1973, I noticed that the male of a copulating pair
of E. pacuvius (Lintner) seemed to squeeze the female’s abdomen repeatedly with
his valvae for several minutes before copulation ended. A copulating pair of E.
persius (Scudder) was then found, and was observed more closely. Mating lasted
38 minutes, and during the last 25 min. at least, the male scraped his left valva

VoLUME 32, NuMBER 4 305

over her convex 7th sternum, by actually bending the valva in the middle such that
the long ventral process and short middle process bent medially, but the base of the
valva and the right valva were relatively stationary.

On a hilltop at Jarre Canyon, Douglas Co., Colorado on 26 May 1978, I observed
a copulating pair of E. persius in which the male scraped his left valva over the
female sternum 7 for 30 min. from capture until both were killed and microscopically
examined while still joined. This pair was examined very closely while in copula.
The male and female abdomens were inclined upward, forming an angle of about
150° ventrally to each other. ‘The male’s uncas fit over the posterior rim of the
female lamella, and beneath the papilla analis. The lower process of the right valva
pressed the membrane above the female sternum 7. The upper process of the right
valva was not visible but probably hooked over the right edge of the lamella. The
upper process of the left valva hooked dorsally over the right edge of the female
sternum 7. The middle and lower process, however, were flexed medially about 30°
over the ventral surface of sternum 7, and were flexed medially another 30° during
each flexion at intervals of about 1 per sec. The female dorsal belt of “scent scales”
was exposed to view during copulation; the ventral hair pencils were also exposed
(but not expanded), and approximately 2 mm of sternum 7 were exposed to view.
With each flexion of the middle and lower processes of the left valva, scraping across
the female stemum 7 occurred, and the lower process of the right valva pressed the
female membrane above sternum 7 inward, while the exposed 2 mm length of
sternum 7 shrank to 1.5 mm as the male’s abdomen telescoped slightly. The male
abdomen exhibited squeezing movements with the valva about 1 per sec. but not
peristaltic movements. When dissected, the female was found to have a full-sized
spermatophore with the usual complement of transparent granules, milky bulb, and
partly formed neck.

Female Erynnis genitalia are somewhat unusual in that sternum 7 is nearly as
heavily sclerotized as the lamella. The lamella is partly telescoped under sternum 7
during copulation, and the male persius scrapes sternum 7 with his left valva over
the bulge on the right side of the asymmetric female lamella. It is noteworthy that
genitalic asymmetry is developed strongly only in subgenus Erynnides and not
strongly in subgenus Erynnis Schrank (Burns 1964, op cit). Most members of sub-
genus Erynnis probably do not exhibit valval scraping because Burns states that
sternum 7 of females is densely scaled in all species except icelus (Scudder & Burgess ).
Asymmetry of genitalia therefore seems associated with asymmetric valval flexion
during copulation. The function of valval scraping is unknown. The hair pencils
were intact after mating of these females and are intact in most museum specimens
which have mated. These observations are reported here because mid-valval flexion
is, to my knowledge, completely unknown in Lepidoptera, and hopefully this note
will stimulate others to help elucidate the function of this strange behavior.

James A. Scott, 60 Estes Street, Lakewood, Colorado 80226.

EDITOR’S NOTE:

While carrying out numerous intra- and inter-specific hand-pairings of Limenitis
during the past 12 years, I have often observed the process of copulation in this
genus. During mating, it is usual for the males to exhibit mid-valval flexion of the
type described above by Dr. Scott. Such observations can be made using a stereo-
microscope without disturbing the mating pair. Both valvae can be seen to flex
inward, perhaps serving to stimulate the female by raking the distal teeth (or hooks )
across her lateral abdominal sternae. The valvae of Limenitis are symmetrical, except
for the placement and number of teeth on the distal tips of the claspers (see Platt,
Frearson, & Graves 1970, Canad. Entomol. 102: 513-533).

AustTIN P. Puatr, Department of Biological Sciences, University of Maryland Balti-
more County, 5401 Wilkens Ave., Catonsville, Maryland 21228.

Journal of the Lepidopterists’ Society
32(4), 1978, 306

CATOCALA ILIA (NOCTUIDAE) FEEDING ON DECAYING FRUIT IN AN
INNER-CITY ENVIRONMENT

The feeding of adult Lepidoptera on over-ripe fruit has been noticed for at least
two centuries, but some recent observations may be of value and interest because
they incorporate data about diurnal activity of a nocturnal moth as well as occurrence
in an inner-city environment.

Occasional accounts have appeared of daytime flight of Catocala, yet adult feeding
of this genus has generally been regarded as crepuscular and nocturnal. Moreover,
little is known about the occurrence of Catocala in cities of some size, although it
appears that the moths may be found in large numbers in such situations; for ex-
ample, I have described Bryant Mather’s collecting of 124 specimens (15 species)
within two days in downtown Jackson, Mississippi (Wilkinson 1971, Michigan
Entomol. 4: 59-60). In seven years of residence on Capitol Hill, Washington, D.C.,
I have found Catocala in abundance, but not as numerous in specimens or species
as in Mather’s situation. Although I have taken so rare a species as C. marmorata
Edw. (daylight, at rest on a wall of the Library of Congress near an entrance lamp,
27 July 1977), the most common species of inner-city Washington is C. ilia (Cramer).

Seated in my enclosed back garden on 22 August 1976, while the sun was shining,
I noticed a large Catocala fly into the grape arbor at 1400. Upon searching, I found
a male C. ilia feeding at a cluster of over-ripe and broken Concord grapes. When
leaving the extensive arbor I flushed another Catocala which was not captured but
was quite probably C. ilia.

In 1977, several ornamental Oriental peach trees in my back garden threw down
their first extensive crop of many hundreds of small peaches. I was unable to attend
to gathering all of these immediately, and a number began to decay. At the same
time I noticed an abnormal frequency of C. ilia on my windows and about my
security floodlight at night. Walking into the garden with a flashlight at 0200 on
2 July I disturbed numerous C. ilia (many of which were taken, identified and re-
leased) feeding on the decaying peaches. With these hints in mind, on 4 July I
kept a long watch in my garden, having purposely left peaches on the ground. At
approximately four hours before dusk I noticed the first ilia flying in the garden.
From that time until dark, C. ilia were sighted in flight at 20 to 40 minute intervals.
I was able to trace perhaps 50% of these individuals, always to or near rotting
peaches. Diurnal and nocturnal feeding continued into early August, until fallen
peaches were no longer edible.

These observations not only suggest further opportunities for study of diurnal flight
and feeding of Catocala, but indicate the desirability of more extensive research into
the occurrence of the genus in urban areas. My data on Catocala in Washington, D.C.
continue to accumulate.

Ronatp S. Wixixinson, The American Museum of Natural History, New York,
New York 10024.

Journal of the Lepidopterists’ Society
32(4), 1978, 307-309

PERIODIC OCCURRENCE OF URANIA FULGENS (URANIIDAE)
IN THE UNITED STATES’

In a recent literature search I came across a one-page article by Leussler (1918,
Entomol. News 29:149) titled, “Interesting butterfly occurrences at Beeville, Texas.”
The first species listed is, “Cydimon poeyi Gundlach—One specimen of this tropical
swallowtail was captured by Miss Pattie Hutchinson at Beeville, June 17, 1916.”
Leussler cited Felder & Felder [1864—1875, Reise der Osterreichischen Fregatte
Novara um die Erde in den Jahren 1857-1859. Zool. Theil 2 (7)], as giving an
illustration of the species which Miss Hutchinson had collected.

Confusion resulted from Leussler’s report, first from the term “butterfly,” and
second from incorrect nomenclature. The oldest valid generic name (D. C. Ferguson,
pers. com.) is Urania Fabricius, 1807, not Cydimon Dalman, 1824. The nomenclature
was further confused by the use of the species name poeyi Gundlach, of Cuban
origin, and superficially close to fulgens according to Gaede [1930, in Seitz, The
Macrolepidoptera of the World, 6:820-831 (Urania). Alfred Kernen Verlag, Stutt-
gart], and only doubtfully distinct (D. C. Ferguson, pers. com.). Thus, what Miss
Hutchinson actually caught was Urania fulgens Walker.

The purpose of this paper is not to make or propose taxonomic changes, but rather
to present the true identity of the specimen recorded by Leussler, and to document
other occurrences of the species within the United States. The specimen which
Miss Hutchinson captured (Fig. 1) is now in the USNMNH together with two
other examples from Texas, one of which is illustrated (Fig. 2). Interestingly,
McDunnough [1938, Mem. So. Calif. Acad. Sci. 1 (1)] did not include this genus
or species in his check list, possibly because Leussler’s report of 20 years earlier
had not come to his attention.

Of the 10 examples of this species collected in the United States over the past
60 years, all except one are from Texas. A migration of this species in Texas also
is recorded. Perhaps there are other existing examples in collections just waiting to
be brought to light. Data on the known examples follow:

FLORIDA: 1 ¢, worn, 9 September 1973, Fort Walton Beach, Okaloosa Co.,
V. J. Farkas (Emmel & Farkas 1974, J. Lepid. Soc. 28:292) (in Kendall collection).
TEXAS: 1 @, worn, 17 June 1916, Beeville, Bee Co., Miss Pattie Hutchinson
(Leussler 1918, cited above) first U.S. record; 1 2, worn, 8 April 1941, Lancaster,
Dallas Co., Mickey Lemmon; 1 @, worn, 11 April 1941, Lancaster, Dallas Co.,
H. A. Freeman (these 3 specimens in the USNMNH); 1 4, worn, 6 April 1941, nr.
San Antonio, Bexar Co., C. O. Neumann (in Kendall collection, ex coll. A. E.
Brower); 1 2, worn, 27 December 1955, College Station, Brazos Co., no coll. label
(in Texas A&M Univ. collection); 1 ? 2, worn, 5 September 1971, George West,
Live Oak Co., John E. Hafernik, Jr., and in his collection; 1 ? ¢, in sealed mount,
W. W. White Elementary School, San Antonio, Bexar Co. (a recent inquiry disclosed
the specimen no longer exists); 2, sex undetermined, 1 in good condition, the other
fair, July 1939, San Benito, Cameron Co., Jack B. Prentiss (given to his high school;
recent inquiry discloses they no longer exist).

Gaede (1930, cited above) gave brief descriptions of the larva and pupa for U.
leilus, but made no mention of a larval foodplant. Beutelspacher (1972, J. Lepid.
Soc. 36:133-137) found a cocoon of U. fulgens in Mexico among the leaves of an
epiphytic bromeliad growing on the trunk of a coconut palm. Smithsonian Institution
Research Reports No. 7, Winter 1974, and Neal G. Smith (pers. com.) confirm
Omphalea diandra L., Euphorbiaceae, as a larval foodplant for U. fulgens; this is a
difficult plant to find because of its climbing habit and foliage production high in
the forest canopy.

Although little has been published on the life history of the uraniids, much has

1 Contribution No. 399. Bureau of Entomology, Division of Plant Industry, Florida Department
of Agriculture and Consumer Services, Gainesville 32602.

308 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

Figs. 1-2. Urania fulgens, females. 1, first United States record, Beeville, Bee
Co., Texas, 17 June 1916, leg. Pattie Hutchinson. 2, Lancaster, Dallas Co., Texas,
H. A. Freeman, 8 April 1941, leg. Mickey Lemmon.

been published on the migratory habits of these moths. Gaede (1930, cited above)
gave general migratory habits of the genus, but indicated that they do not reach
the United States; no doubt Leussler’s report had not come to his attention. Williams
(1937, Nat. Geog. Mag. 71:568-585) gave Mexico, Nicaragua, Costa Rica, and
south to Ecuador as the migratory area. Valerio G. (1966, Univ. Costa Rica journal,
Cratera 1:40-45) gave the period, magnitude, direction, and speed of flight, and
mentioned an attraction to shiny objects. He indicated also that U. fulgens migrated
in Veracruz, Mexico; Atlantic coast of Honduras, Rivas isthmus of Nicaragua, Panama,
and Bogota area of Colombia. He speculated that the larvae may feed on tree-top
foliage, thus avoiding detection by collectors. Young (1970, J. N. Y. Entomol. Soc.
78:60-70) documented the daily flight activities of U. fulgens in Costa Rica where
an observed migration lasted 42 days; the life expectancy was estimated to be 28
days for males and 34 days for females. Smith (1972, Carib. J. Sci. 12:45-58)
documented the population fluctuations and migrations of Urania moths in lower
Central America, the Lesser Antilles, and in northern South America since 1868.
He found no clear cut periodicity of movement, but the most frequent interval was
8 years.

In July 1939 at San Benito, Cameron Co., Texas, Jack B. Prentiss observed a
migration of U. fulgens during the morning hours, in a pasture behind his home.
In personal communication he stated, “The flight was rather extensive; there were
always a few in sight for the better part of 4 hours. They were all flying in a
due north direction. During the height of the flight they were rather numerous;
20 to 30 could be seen at one time. Most were flying too high to be netted and
all were flying fast. Those flying low were most elusive and one had but a single
chance to net them. During the course of the flight I did manage to take 2 speci-
mens; one was in extremely good condition, the other only fair. Both specimens
were placed in the high school collection. I have since checked to see if they were
still there but found they have long since been discarded.”

ACKNOWLEDGMENTS

or literature citations I am indebted to Cyril F. dos Passos, William D. Field,
Thomas C. Emmel, and Kent H. Wilson. For specimens or for field information on

VoLUME 32, NuMBER 4 309

specimens, I wish to thank Thomas C. Emmel, John E. Hafernik, Jr., Richard S.
Peigler, Jack B. Prentiss, and A. E. Brower. To Douglas C. Ferguson, I am espe-
cially grateful for providing not only data on the Texas examples in the National
Collection, but also for the photographs used in this paper, for confirming the
determination of the Florida specimen, and for reviewing the manuscript.

Roy O. KENDALL, Route 4, Box 104-EB, San Antonio, Texas 78228.”

> Research Associate, Florida State Collection of Arthropods, Division of Plant Indusiry, Florida
Department of Agriculture and Consumer Services.

Journal of the Lepidopterists’ Society
32(4), 1978, 309

A PARTIALLY ALBINIC ABERRATION OF PHYCIODES THAROS
(NYMPHALIDAE )

On 25 July 1977, I took a partially albinic male aberration of Phyciodes tharos
Drury (Fig. 1) in Upper Tyrone Township, Fayette Co., Pennsylvania, at an eleva-
tion of 1100 ft (335 m). All the normally tawny or brown coloration characteristic
of this species was replaced by an extremely pale, orange-tinged cream color. The
black markings were not affected. I know of no similar specimens. The aberration
has been deposited in the collection of the Peabody Museum of Natural History,
Yale University, New Haven, Connecticut.

Cuar.es G. Ottver, R. D. 1, Box 78, Scottdale, Pennsylvania 15683.

Fig. 1. Phyciodes tharos Drury: A., B. pale (partially albinic) male aberration,
dorsal and ventral sides; C., D. typical male from same locality, dorsal and ventral.

Journal of the Lepidopterists’ Society
32(4), 1978, 310

A RECORD OF AGRIAS AMYDON (NYMPHALIDAE) FROM COSTA RICA

Two female specimens of Agrias amydon Hew. subspecies (?) were taken in Parque
Santa Rosa, Guanacaste Province, Costa Rica. This is the first substantiated record
of this genus from Costa Rica. Agrias zenodorus smalli Miller & Nicolay has been
anticipated from Costa Rica and there exists in the literature a supposed sight record
from Turrialba, Cartago Province, Costa Rica (Miller & Nicolay 1971. Bull. Allyn
Mus. (1):1-5). It was totally unexpected to find Agrias in a habitat like Guanacaste
where there is a strongly marked dry season in contrast to the usual wet forest habitats
in other countries where the genus Agrias occurs.

I have compared the specimens with A. amydon in the U.S. National Museum,
the Carnegie Museum, and the Allyn Museum of Entomology and have found them
to be differently marked on the dorsal surface of both fore and hind wings than the
comparative material. Description of this A. Amydon subspecies is impossible on the
basis of only two female specimens and must await more material.

On the basis of the specimens examined in the above mentioned museums and in
the literature available to me, A. amydon is recorded only as far north as Colombia.
These specimens represent a considerable range extension and a new record for
Costa Rica. One specimen has been placed in the collection of the Allyn Museum
of Entomology, Sarasota, Florida.

Pump J. DeVries, Museo Nacional de Costa Rica, Department de Historia Natural,
Apartado 749, San Jose, Costa Rica.

Fig. 1. Agrias amydon Hew., subspecies (?), female: dorsal (left), ventral

(right). Wingspan = 6 mm.

Journal of the Lepidopterists’ Society
32(4), 1978, 310-311

ADDITIONAL FUNCTION OF THE LEPIDOPTERAN PROBOSCIS

The most unique part of the lepidopteran body form is the proboscis. This pre-
hensile tube functions mainly as a suction device for nutrient procurement. A second,
but minor, function has been observed in a variety of unrelated butterflies and moths

VoLUME 32, NUMBER 4 311

—fluid “pumping.” Reports describe these individuals as having an extended pro-
boscis at a moisture source and simultaneously voiding fluid from the anus (Clench
1957, Lepid. News 11:18-21; Roever 1964, J. Res. Lepid. 3:103-120; Hessel 1966,
J. Lepid. Soc. 20:242; Jobe 1977, Entomol. Gaz. 28:8). Interpretations of this be-
havior have been speculative generalizations. Personal observations have revealed a
third—also minor—function of the lepidopteran proboscis.

A female Atalopedes campestris (Boisduval) (Hesperiidae) was found floating with
fluttering wings on the water surface of a wading pool in a residential backyard in
Austin, Travis Co., Texas. The skipper did not appear to be severely injured, probably
due to a relatively short period of partial submergence. Nevertheless, the scales
were quite wet.

Subsequent observation of this individual revealed that it was rapidly probing the
scaly covering of the anterior part of its body by continued manipulation of its
proboscis. Most of the probing involved ventral and lateral scales of the thorax. A
definite color-lightening effect was observed. This change in coloration would indi-
cate a reduction in the amount of water which had a plastering effect upon the
scales.

Decrease in amounts of water present among the scales could result from two
effects of probing by the proboscis. Simple separation of adjacent matted scales by
mechanical movement of the proboscis would increase evaporation rates because of
an increase in scale surface area exposed to air. Also, water could be and probably
was being physically removed by suction via the proboscis.

The primary behavioral regime enlisted in this task would involve an activity
related to previous function—suction action of the proboscis. Therefore, physical
removal of water should be regarded as the initial function upon which natural
selection acted. Enhancement of evaporation rate by scale separation initially was an
ancillary result of this behavior. Such enhancement could be further increased by
selection favoring rapid random movements as opposed to sucking up water at one
spot and relying on capillary pressure to maintain a continuous film of water.

Although this behavior was observed to function to remove water from scales
following partial immersion in water, it could also function to remove moisture from
rain (dew?) on butterflies roosting in exposed sites. Removal of water may be
important for several reasons. Flying ability may be reduced if water is present in
sufficient amounts to appreciably increase the weight of the insect. A matted water/
scale film present on the body surface may interfere with spiracular inhalation or
favor development of pathogenic populations. Even if most air were inhaled via
abdominal spiracles, removal of thoracic surface water may decrease abdominal and
wing surface water via capillary action.

RayMonp W. Neck, Pesquezo Museum of Natural History, 6803 Esther, Austin,
Texas 78752.

JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

INDEX TO VOLUME 32

(New names in boldface )

aberrations, 221, 309
Abraxas grossularia, 233
Achalarus lyciades, 201
Achlyodes thrasa tamenund, 112
Acraea encedon, 233

A. lapitha, 266

A. subhyalina, 268
Acraeidae, 233
Acraeinae, 261
Actinote calderoni, 261, 264

A. lapitha zilchi, 261, 266

A. stratonice oaxaea, 261

A. subhyalina, 268

A. thalia, 261
Adelpha, 171
Aeaea venifica, 235
Aganisthos, 171
Agathymus, 107
Ageronia, 169
Agonopterix dimorphaella, 235
Agraulis vanillae, 178, 211
Agrias, 169, 310

A. amydon, 310

A. zenodorus smalli, 310
Aiello, Annette, 135
Amaryssus astenous, 116

A. minos, 116
Amblyscirtes aesculapius, 28

A. samoset, 28, 200

A. vialis, 200
Amphichlora, 169
Amplypterus gannascus, 56
Anaea andria andria, 200, 211

A. eurypile, 172

A. morvus, 172

A. pithyusa, 172
Anartia amathea, 135

A. fatima, 135, 172

A. jatrophae, 172
Ancyloxypha numitor, 146, 201
Anthocharis midea, 202
Antithesia montana, 251

A. sara, 226
Apanteles, 109, 128
Apatura, 171
Apaturidae, 212
Apodemia palmeri, 209
Archeprepona, 171
Argynnis paphia, 63, 146
Asterocampa celtis celtis, 28, 203

A. clyton clyton, 203

A. leilia, 209

Atalopedes campestris, 200, 311
Atopothoures ovaliger, 55
Atrytone delaware delaware, 200
Atrytonopsis hiana hiana, 200
Auctor hujus operis, 116
Austin, George T., 207
Autochton cellus, 28, 201
Baker, James H., 241
Balogh, George, 289
Bandera, 105
Barcita, 244
Bartlett-Calvert, Wm., 251
Batrachedra illusor, 235
Battus philenor, 28, 157, 201, 209
behavior, 38, 56, 233, 282
Berenbaum, May, 303
Biblis, 171
B. hyperia aganisa, 113
bionomics, 107
Blanchard, André, 55, 103
Blau, William S., 138
Blood-spot skipper, 107
Boloria bellona, 28, 204
B. selene myrina, 204
book reviews, 63, 144, 239
Botys furnaclis, 130
Bowers, Deane, 140, 282
Braconidae, 128
Brephidium exilis, 211
Brewer, J. Wayne, 123
Butler, Linda, 198
Calephelis borealis, 202
C. muticum, 142, 200
Calpodes ethlius, 108
Calydna lusca, 46
C. virginiensis virginiensis, 202
Callaghan, Curtis j., 37
Callizona, 169
Callophrys, 3, 143
C. augustinus croesiodes, 202
C. barryi, 4
C. byrnei, 4
C. gryneus, 3
C. g. castalis, 7
C. g. gryneus, 6, 202
C. g. sweadneri, 6
C. henrici henrici, 24, 28, 202
C. hesseli, 3
C. irus irus, 200
Grlokiees
C. nelsoni, 3
C. niphon niphon, 202

VoLUME 32, NUMBER 4

C. polios polios, 202
@.4siva, 3
C. s. juniperaria, 6, 14
C. s. mansfieldi, 14
C. turkingtoni, 3
Callosamia angulifera, 191
C. promethea, 59, 191, 233
C. securifera, 194
Calpodes ethlius, 108, 200
Calycopis cecrops, 202, 236
Calydna lusca, 46
Castniidae, 139
Catocala, 63
. grynea, 222
. habilis, 222
. ilia, 306
. marmorata, 306
. micronympha, 221
. muliercula, 222
. unijuga, 222
Catagramma, 171
Catonephele, 170
C. numilia, 172
C. nyctimus, 172
Catonephelini, 160
Celastrina argiolus, 34
C. a. bakeri, 240
C. ebenina, 20, 203
C. pseudargiolus, 20, 28, 203, 226
Cercyonis meadii, 146
C. pegala pegala, 204
Charaxinae, 170
Charaxini, 160
Chedra inquisitor, 235
Chelonia midas, 273
Chew, Frances S., 129, 159
Chionodes asema, 235
Chlosyne gorgone carlota, 203
C. harrisi harrisi, 200
C. janais, 59
C. lacinia crocale, 211
C. nycteis nycteis, 203
Chrysocharis, 128
Glark J. iF. Gates, 251
Clench, Harry K., 273, 277
climatic regimes, 111
Coea acheronta, 170
Cogia hippalus, 209
Coleotechnites, 118
. edulicola, 118, 123
. milleri, 123
. moreonella, 120
. ponderosae, 118
. ponderosana, 122
. starki, 123
Colias eurytheme, 28, 202, 209, 228

aS SiGie@

PD} GQ) ©} >)

C. cesonia, 113, 200, 209

CC intenor 43 202,

C. philodice, 28, 202
collection, 235
Colobura dirce, 160
Coloburini, 160, 170
Coloradia, 97
Copacodes aurantiaca, 211
courtship, 41
Cosmopterigidae, 235
Covell, Charles V., Jr., 221
Cryptotainia canadensis, 237, 304
Cyclogramma, 171
Cydimon poeyi, 307
Cyllopsis dospassosi, 75

C. gemma freemani, 84

C. nayarit, 84

C. pephredo, 84

C. pseudopephredo, 84

C. wellingi, 75
Cynthia, 94

C. annabella, 95

C. cardui, 95, 203

C. carye, 95

C. virginiensis, 95, 203
Danaidae, 170, 204
Danainae, 116
Danaus gilippus, 209

D. plexippus, 204, 209
Demons, IRs ling lal 2s)
descriptions, 75, 175, 261
DeVries, Philip J., 310
Diaethria, 171
Dichomeris glenni, 235
Dicladocerus, 128
Dimock, Thomas E., 88
Drees, Bastiaan M., 198
Dryas julia moderata, 111
Dymasia chara, 209
Dynamine dyonis, 111
Eberlie, W. J. D., 226
Electrostrymon angelia angelia, 139

E. a. boyeri, 140

E. endymion, 139
Endothenia microptera, 235
Epargyreus clarus clarus, 28, 201
Epiblema naomi, 235
Epimetes, 116
Epinotia, 256

E. atristriga, 235
Epiphile, 170
Erora, 279

E. laeta, 35, 140, 202
Erunniss 24354 6 305

E. baptisiae, 201

313

314 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

E. brizo brizo, 201 F. thalia, 279
E. b. lacustra, 28, 61 F. w-album, 277

E. funeralis, 209 foodplant, 3, 20, 86, 236, 303

E. horatius, 201 Gelechiidae, 118, 123, 235

E. icelus, 201 genitalia, 7, 304

E. juvenalis, 25, 28, 201 genetic factor, 231

E. lucilius lucilius, 201 geographic distribution, 3, 259, 291

E. martialis, 201 Geometridae, 224

E. pacuvius, 305 Gilbert, Lawrence E., 54

E. persius, 200, 305 Glaucopsyche lygdamus nittanyensis, 203

E. zarucco zarucco, 200 Godfrey, George L., 235
Euchloe olympia olympia, 202 Goya ovaliger, 55
Eucosma uta, 235 G. stictella, bo
Eucosminae, 256 Graphium marcellus, 201
Eueides cleobaea zorcaon, 113 Gynaecia, 169
Eulophidae, 128 Gynaeciidae, 170
Eulythis mellinata, 224 Gynaeciini, 160
Eunica, 169 gynandromorph, 138
Euphaedra ceres, 116 Gypsonoma, 256
Euphydryas anicia bakeri, 240 habitat, 20

E. chalcedona, 226 habitss 1235 sie

E. phaeton, 204, 282 hairstreak, 277
Euphyes bimacula, 200 Hamadryas, 171

E. dion dion, 200 H. amphinome, 172

E. conspicua conspicua, 200 H. februa, 172

E. vestris metacomet, 200 H. guatemalena, 172
Euptoieta, 169 Hamadryadini, 160

E. claudia, 204, 209 Hardwick, D. F., 49
Euptychia areolata areolata, 200 Harkenclenus titus mopsus, 202

E. cymela cymela, 28, 904 Heliconiidae, Jia 170, 185, 212

E. gemma gemma, 204 Heliconius charitonius, 172

E. hermes, 28 H. c. vasquezae, 111

E. h. sosybius, 204 H. erato petiverana, 44

E. rubricata, 146 H. ethilla, 43
Euptychiini, 75 H. petiveranus, 172
Eurema, 39 Heliothis fastidiosa, 49

E. lisa, 178, 202 H- glomosa) BO

E. nicippee, 202, 209 H. imperspicua, 51
Euristrymon, 277 H. inclara, 51

E. ontario ontario, 200 H. lanul, 51
Everes amyntula, 209 He Ree a2

Y ,

E. comyntas, 34, 146, 203 EGE, oe
ay, Oe tee Lares H. rubiginosa, 52
Exartema comandranum, 235 H. siren, 53
Felderia eximium, 280 H. spectanda, 53
Feniseca tarquinius tarquinius, 202 H. sulmala, 53
Fiji, 130 Hermiargus ceraunus, 209
Finke, Mark, 236 H. isola, 209
lixsenia acaciae, 277 Hemileuca chinatiensis, 97

F. favonius, 279 H. diana, 99

F. herzi, 277, 279 H. electra, 99

F. ilicis, Bag H. griffini, 97

F’. ontario, 279 H. grotei, 99

F. pruni, 277, 279 H. hera hera, 59, 99

F. polingi, 279 Hemimene, 258

F. spini, 277 Hesperia, 146

VOLUME 32, NUMBER 4

. attalus attalus, 200

. leonardus, 201

. metea, 200

. Sassacus sassacus, 201
. columbia, 61

me

Hesperiidae, 107, 112, 143, 199, 225,

304, 310

Hesperiinae, 185, 189
Historis odius, 160
Hodges, Ronald W., 118
Homoptera aemona, 242, 243

H. discisigna, 243

H. peruncta, 242

H. ustipennis, 242
Hyalophora, 191

H. cecropia, 194, 233

H. columbia, 143

H. gloveri gloveri, 233
hybridization, 95, 191, 226, 289
hybrids, 191, 226, 289
Hylephila phyleus, 189, 201, 211
Hymenoclea salsola, 215
Hymenoptera, 128
Hypercallia, 251
Hypolimnadidi, 170
Hypolimnas misippus, 170, 233
NG-Z.N., AS
Incisalia iroides, 44, 146
insectivorous plant, 129
interactions, 65
inter-specific hybrids, 191, 226, 289

“arthechippus,” 289

eurymedon X rutulus, 226

lorquini < archippus, 289

“rubidus,” 289

“weidechippus,” 289
Ithomiidae, 170
Jennings, Daniel T., 123
Johnson, Kurt, 3
Junonia coenia, 203
Kendall, Roy O., 75, 86, 307
Kohler, Steve, 1, 57
Kricogonia lyside, 59
Laetilia, 105
Lamas. Gerardo, 116
Larisa subsolona, 256
Laspeyresia, 256
Laspeyresiinae, 256
lectotype designations, 49
Lederhouse, Robert C., 145
Leptotes marina, 209
Lerema accius, 178, 200
Lerodea eufala, 200, 211
Lethe anthedon, 28, 204

L. appalachia, 204

L. creola, 200

315

L. eurydice eurydice, 204

L. europa malaya, 64
Letis incipiens, 242
letter to the editor, 54
Leucochimona philemon, 46
Libytheana bachmanii bachmanii, 203

L. b. larvata, 209

Libytheidae, 203, 212
life cycle, 88, 160, 282
life histories, 123
Limenitidini, 170
Limentis, 228, 289, 305
L. archippus, 203, 228, 289

L. a. floridensis, 289

L. arthemis arizonensis, 203

L. a. arthemis, 203, 289

ik, Ga, asvimmabs, 2S, sie 208) CAE
228, 289

L. lorquini, 289

L. weidemeyerii, 289

Lycaeides melissa, 59

L. m. samuelis, 200

Lycaena arota, 44
L. hyllus, 202
L. phlaeas, 146, 202
Lycaenidae, 59, 139, 140, 146, 199, 212,

236, 277

Lycorea pieteri, 116
malaise traps, 178
Marpesia, 171

M. berania, 43
McAlpine, Wilbur S., 142
Meander felsina, 37
Mecyna furnacalis, 130
Megistanis, 170
Mellichamp, T. Lawrence, 20
Mestra, 171

M. amymone, 112
Metria, 242
Mexican Actinote, 261
Meyrick, 130, 251
microlepidoptera, 235
mid-valval flexion, 304
migration, 178
Miller, Jacqueline Y., 261
Miller, Lee D., 63, 64, 75, 139, 261
Miller, Thomas A., 233
Miller, William E., 256
mimicry, 157, 170
Ministrymon leda, 209
Mitoura, 3
Momphidae, 235
Morphidae, 65
Morpho achilles, 73

M. amathonte, 73

MENCUDTISt

M. granadensis, 66, 73

316 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

M. peleides, 65 P. a. temenes, 273
M. theseus, 73 P. blomfildia, 169
mosaics, 228 P. ceres, 116
Munroe, Eugene, 130 P. cresphontes cresphontes, 201, 211
Mutuura, Akira, 130 P. demodocus, 146
Muyshondt, Alberto, 160 P. eurymedon, 226
Muyshondt, Albert, Jr., 160 P. glaucus, 28, 138; lav) 201226
Najas ceres, 116 P. joanae, 237, 304
Nastra lherminier, 201 P. multicaudatus, 226
Nathalis iole, 200, 209 P. polyxenes asterias, 138, 145, 157,
natural history, 88 201, 228, 236, 303
Neck, Raymond W., 107, 111, 224, 310 P. rutulus, 226
Neil, Kenneth, 224 P. troilus, 28, 157, 201
Neilsen, M. C., 142 P. xuthus, 228
Neolycaena sinensis, 280 P. zelicaon, 226
new foodplant, 236, 303 Papilionidae, 1, 138, 139, 145, 226, 236,
new genera, 103, 116 Dns, BIO}
new generic assignment, 130 Parnassius clodius gallatinus, 1
new record, 236 Peigler, Richard, S., 191
new species, 75, 78, 82, 97, 118, 256 Peridroma plecta, 134
new subspecies, 75, 248, 261, 273 Peridromia, 169
Nica, 171 Perimede maniola, 235
Noctuidae, 134, 241, 306 Periploca cata, 235
Nordmannia, 277 Peteroma, 244
notes and news, 19, 48, 159 phenology, 207

Nymphalidae, 59, 88, 116, 135, 146, 160, Phocides lilea sanguinea, 107
185, 199)919) 924031) 9e8 961, Pl nolbus Ifeau lon

282, 289, 309, 310 P. pygmalion okeechobee, 108
Nymphalini, 170 Phoebus agarithe maxima, 59
Nymphalis antiopa, 203, 282 P. sennae, 178, 209

N. milberti milberti, 203 P. s. eubule, 202

N. vau-album, 200, 282 P. philea, 223
Nymphidium lisimon attenuatum, 46 Pholisora catullus, 201, 209

N. mantus, 46 Phyciodes batesti, 203
obituaries, 142, 240 P. tharos, 28, 146, 203, 231, 309
Oecophoridae, 235, 251 Pieridae, 59, 129, 139, 170, 185, 200, 212,
Oeneis alberta, 57 DPN), ONS,

O. chryxus strigulosa, 143 Pieris brassicae, 139
Ogunwolu, E. O., 175 P. napi oleracea, 129, 226
Oidaematophorus glenni, 235 P. protodice, 202, 209, 228
Olethreutidae, 235, 256 P. rapae, 202, 230
Oliver, Charles G., 231, 309 P. virginiensis, 28, 35, 202
Ollia parvella, 103 pine feeding species, 118
orange bands, 135 Platt, Austin P., 289, 305
Ornithoptera priamus, 138 Poanes hobomok, 28, 200

O. victoriae, 138 P. massasoit hughi, 200
Ostrinia, 130 P. m. massasoit, 200
oviposition, 233, 236 P. viator, 200
Pammene, 258 P. yehl, 200
Panoquina ocola, 178, 200 je zabulon, 200
Panthiades m-album, 200 Pogue, Michael G., 236
Papilio andraemon, 273 Polites coras, 201

P. aristodemus, 273
P. a. bjorndalae, 275
P. a. driophilus, 273
P. a. ponceanus, 273

. mystic, 201

. origenes origenes, 201
. themistocles, 201

. vibex vibex, 200

ae Bene pias ac!

VoLUME 32, NUMBER 4

Polychrosis sambuci, 235
Polydesma, 243
Polygonia comma, 28, 203
P. faunus faunus, 203
P. interrogationis, 28, 203
P. progne, 203
Pompeius verna verna, 200
population structure, 37
Precis coenia, 178, 224
predation, 134
Prepona, 170
proboscis, 310
Problema byssus, 200
Proeulia, 253
P. montana, 253
Pseudohazis, 97
Pseudonica, 170
Pteromalidae, 128
Pterophoridae, 235
Pycina, 170
Byralidae,, po, 103, 175
Pyrausta homaloxantha, 130
Pyraustinae, 130
Pyrginae, 184

Pyrgus centaureae wyandot, 201

P. communis, 201, 209

Pyrrhogyra, 169

Rawson, George W., 289
rediscription, 103
Restinga butterflies, 37
Rhododipsa aden, 49
Rhopalocera, 111, 198
Riodinidae, 37, 200
Robb, Jeff, 56, 59
roosting, 145
Rostralaetilia, 103
Rawlins, John Edward, 146
Safia, 242

Samia cynthia, 191
Satummidae, 59, 97, 191, 233
Satyridae, 57, 86, 200, 204
Satyrium acaciae, 273, 280
. acadica acadica, 200
. acadicum, 281
. adenostomatis, 281
. auretorum, 281
. behrii, 280
. calanus, 281
. californicum, 281
Sue. jalicen 202
. caryaevorus, 200, 281
. edwardsii, 202, 281
. eximium, 281

. fuliginosum, 280
- tlicis, 281

NN

RNNRNNN

RNNNN

. kingi, 281
. latior, 281
. liparops, 281
S. l. strigosa, 202
. lunulatum, 280
. myrtale, 280
. pretiosum, 281
. saepium, 281
. sassanides, 280
. sinensis, 281
. spini, 280
. sylvinum, 281
. tetra, 281
. tengstroemii, 281
. w-album, 280
Sceliodes cordalis, 177
S. laisalis, 175
Schinia approximata, 49
S. Gr, DO
S. dolosa, 49
. hanga, 50
. labe, 49
. lora, 49
. neglecta, 52
. obscurata, 52
. pyraliodes, 49
. tanena, 53
ultima, 53
Scott, James A., 304
Scriber, Mark J., 236
Sebethis, 116
Shapiro, Arthur M., 223, 228
Shields, Oakley, 61
Silberglied, Robert E., 135
Siproeta epaphus, 43, 172
S. stelenes, 172
Smith, Dwight G., 134
Smyrna bella, 169
S. blomfildia, 160
S. karwinskii, 160
S. pluto, 169
Sorhagenia baucidis, 235
Speyeria aphrodite, 146, 204
S. atlantis atlantis, 204
S. cybele cybele, 28, 204
S. diana, 204
Sphingidae, 56, 139
species diversity, 218

NNN

NNNNNRNRNRNHnRNHNHNN

ANNNNnNNNN

Splendeuptychia kendalli, 75, 86

S. salvini, 78
Staphylus ceos, 209
S. mazans hayhurstii, 201
Stevens, Robert E., 118, 123
Strymon, 278
S. melinus, 209
S. m. humili, 202

S17

318

Strymonidia thalia, 277
Systematics, 1, 49, 75, 97, 103, 116, 118,
130: 139. 241 25125652615 2738;
TT
Tampa, 105
Taygetis mermeria excavata, 75
T. m. griseomarginata, 75
Teladoma incana, 235
Temenis, 170
temporal distribution, 86
Thecliolia, 280
Thymelicus lineola, 146, 200, 225
Texola eulauda perse, 211
Thorybes bathyllus, 201
T. confusis, 200
T. pylades, 28, 201
Tilden, J. W., 241
Tmolus azia, 236
Todd) Bok. 244
Tortricidae, 251
Trogus pennator, 303
Tuskes, Paul M., 97
unusual occurrences, 111
Urania fulgens, 307
U. leilus, 307
Urbanus dorantes, 178
U. proteus, 178, 200
Uresiphita, 130
Urticastrum mexicanum, 166

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Vanessa annabella, 88, 211
V. atalanta, 203, 211
V. a. rubria, 93
V. cardui, 93, 209
V. virginensis, 93
voltinism, 216
Wagner, Warren H., Jr., 20, 226
Walker, Thomas J., 178
Wallengrenia egeremet, 201
Walshiidae, 235
Welderella, 103
W. parvella, 105
Wilkinson, Ronald S., 306
Xanthopsamma, 130
X. aurantialis, 132
X. homaloxantha, 132
X. youboialis, 132
Xylis setipes, 242
Young, Allen M., 65
Zagrammosoma multilineatum, 128
Zale discisigna discisigna, 241
Z. notipennis, 241
Z. peruncta, 241
Z. p. ineipiens, 248
Z. setipes, 241
Z. s. £. postmedialis, 246
Zaretis, 171
Zerene cesonia, 113, 209
Zinnia pumila, 215

PLANT INDEX FOR VOLUME 32

Acacia greggii, 209
Acer rubrum, 33
A. saccharum, 33
Adiantum pedatum, 34
Aegopodium podograria, 237
Aesculus octandra, 33
Ailanthus altissima, 192
Althaea rosea, 92
Anethum graveolens, 303
Angelica atropurpurea, 237, 303
Antirrhinum, 224
Arachis hypogea, 67
Aruncus dioius, 20, 26
Asclepias syriaca, 155
Aster novae-angliae, 154
A. simplex, 23]
Asteraceae, 32
Astilbe biternatum, 32
Athyrium filix-femina, 34
Bambusa aculeata, 79, 86
Betula lenta, 33

Bidens pilosa, 185
Botrychium virginianum, 34
Callistemon citrinus, 109
Canna, 108
Caprifoliaceae, 29, 285
Capsicum annuum, 175
C. frutescens, 175
Carex plantaginea, 34
Carpinus caroliniana, 33
Celtis pallida, 207
Cercidium microphyllum, 207
Cercocarpa betuloides, 99
Chamaecyparis thyoides, 3, 7
Chelone glabra, 282
Cicuta bulbifera, 237, 303
C. maculata, 303
Cimicifuga racemosa, 34
Coleogyne ramosissima, 97
Conium maculatum, 237, 304
Cornaceae, 29

VoLUME 32, NUMBER 4

Cornus florida, 33
C. racemosa, 29
C. stolonifera, 29

Cryptotaenia canadensis, 237, 303

Cupressaceae, 3
Cynodon dactylon, 86
Dactylis glomerata, 151, 155
Dacus carota, 237, 303
Dioclea, 72

D. wilsoni, 67
Diplacus, 224
Drosera rotundifolia, 129
Duranta repens, 109
Erythrina crista-galli, 67
Euphorbiaceae, 307
Fagus grandifolia, 33
Gramineae, 86
Geranium maculatum, 24
Heracleum maximum, 237
Hydrangea arborescens, 21
Hymenoclea salsola, 215
Impatiens capensis, 34
Inga, 72
Ipomoea coccinea, 215
Juniperus, 3

J. ashei, 4

J. californica, 6

J. communis, 6

J. depeana, 4

J. flaccida, 7

J. horizontalis, 4
. monosperma, 4
. occidentalis, 4
. osteosperma, 4
. pinchotii, 4
. scopulorum, 5
. silicicola, 6
. virginiana, 4
Kickxia spuria, 224
Laportea canadensis, 34
Leguminosae, 65
Levisticum officinale, 238
Lindera benzoin, 33
Linaria, 224

L. vulgaris, 151
Lippia, 224

Cy ay ay ey ey ey

Liriodendron tulipifera, 34, 192

Lonicera japonica, 286
L. morrowi, 151

Lycopersicum esculentum, 175
Machaerium floribundum, 65, 72

Magnolia acuminata, 236
Malwa, 92

M. parviflora, 92
Malvaceae, 92

Marcgraviaceae, 45
Medicago sativa, 67
Mirabilis multiflora, 215
Moraceae, 166
Mucuna, 72
Myrtaceae, 107
Norantea brasiliensis, 45
Omphalea diandra, 307
Opuntia fulgida, 209

O. spinosior, 209

Osmorhiza claytoni, 237, 303

O. longistylis, 237, 303
Oxypolis filiformis, 303
Passifloraceae, 111
Passiflora lutea, 111
Pastimaca sativa, 237
Penstemon azureus, 224
Phleum pratense, 151, 155
Pinus contorta, 123

P. edulis, 123

P. jeffreyi, 123
Plantaginaceae, 224
Plantago, 224
Potentilla recta, 151, 153
Prosopis juliflora, 207
Prunus serotina, 233, 290
Psidium cattleianum, 107

P. guajava, 107
Ptilimnium capillaceum, 303
Quercus dumosa, 61

QO. durata, 61
Rhododendron, 36
Ribes alpinum, 224

R. nigrum, 224

R. rubrum, 224
Rutaceae, 236
Salix babylonica, 290
Sanicula gregaria, 237
Saxifragaceae, 32
Scrophulariaceae, 224
Sida, 92
Sidalcea malvaeflora, 92
Sium suave, 303
Solanaceae, 175
Solanum macrocarpon, 175

S. melongena, 175
Solidago altissima, 151, 154
Sphaeralcea ambigua, 92
Spermolepis divaricata, 303
Spiracea, 28

Taenidia integerrima, 237, 303

Thaspium barbinode, 304
Thuja occidentalis, 129
Tiarella cordifolia, 34
Tilia americana, 34

Tragopodon pratensis, 32, 151, 153

319

320 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Trillium, 34 Urtica holosericea, 90

Umbelliferae, 236 U. urens, 92

Urera baccifera, 166 Urticastrum mexicanum, 166
U. caracasana, 166 Verbenaceae, 109, 224

Urticaceae, 92, 160 Vibernum lentago, 29

Zinnia pumila, 215

ERRATA

p. 20, abstract, line 7, W. ebenina = C. ebenina

p. 35, third line from bottom, Marylan = Maryland

p. 134, note title & Vol. 2 Contents on back cover, Peridoma = Peridroma

p. 172, line 2, V. epaphus = S. epaphus

p. 214, third paragraph, line 7, C. caesonia = C. cesonia

p. 224, second paragraph, line 4, E. mellinate = E. mellinata

Vol. 3 Contents on back cover, fourth line from bottom, Mark Fink = Mark Finke

(The Editor regrets these errors )

Date of Issue (Vol. 32, No. 4): 28 February 1979

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Us. &

CONTENTS

Tue ZALE SETIPES SPECIES COMPLEX (LEPIDOPTERA: NOCTUIDAE).
Beh. Todd 20000 Se

A Lost AND MispLACeD TAXON (LEPIDOPTERA: TORTRICIDAE). J. F.
Gates Glarke 2.0 +0

LARISA SUBSOLANA, A New GENUS AND SPECIES OF MOTH FROM
EASTERN NortH AMERICA (OLETHREUTIDAE). William E. Miller

Notes ON MEXICAN ACTINOTE (NYMPHALIDAE: ACRAEINAE) AND THEIR
RELATIVES, WITH DESCRIPTION OF A NEW SupssPEciEs. Jacqueline
Y. Miller & Lee D. Miller 0

PAPILIO ARISTODEMUS (PAPILIONIDAE) IN THE BAHAMAS. Harry K.
Clench

THe NAMEs OF CERTAIN HoLarctic HAIRSTREAK GENERA (LYCAE-
NIDAE). Harry K..Clench 0 ee

OVER-WINTERING BEHAVIOR IN EUPHYDRY AS PHAETON (NYMPHALIDAE).
M. Deane Bowers

INTER-SPECIFIC HYBRIDIZATION INVOLVING LIMENITIS ARCHIPPUS AND
irs CONGENERIC SPECIES (NYMPHALIDAE). Austin P. Platt, George
W. Rawson, & George Balogh

GENERAL NOTES

Taenidia integerrima, a new foodplant record for Papilio polyxenes
(Papilionidae).. M. Berenbaum ‘22.0000
Mid-valval flexion in the left valva of asymmetric genitalia of Erynnis
(Hesperiidae). James A. Scott (i000
Catocala ilia (Noctuidae) feeding on decaying fruit in an inner-city en-
vironment. Ronald S, Wilkinson. 0.0.0.0.
Periodic occurrence of Urania fulgens (Uraniidae) in the United States.
Roy O.. Kendall 0000000008 i OS
A merely albinic aberration of Phyciodes tharos (Nymphalidae). Charles
G. Oliver sl a 0 A
A record of Agrias amydon (Nymphalidae) from Costa Rica. Philip J.
DOV 148 esc ces ID Oa I A

Additional function of the Lepidopteran proboscis. Raymond W. Neck -..

Eprror’s Nore

a a a ek be St me Fb en tr ts eo ef oh a Se Pr mS

241

251

256

261

273

277

282

289

Volume 33 1979 Number 1

JOURNAL

of the

LEPIDOPTERISTS’ SOCIETY

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cat 4 June 1979

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Cover illustration: Third instar larva of Limenitis archippus Cramer (Nymphalidae)
preparing to enter winter diapause. The larva is resting on the lip of its hibernaculum
constructed from the basal portion of a chewed tubular willow leaf (Salix babylonica
Linnaeus) covered with silk. In the autumn such larvae begin facultative diapause in
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JOURNAL OF

Tue LeprpopreRiIsts’ SOCIETY

Volume 33 1979 Number 1

Journal of the Lepidopterists’ Society
33(1), 1979, 1-5
PAITITIA NEGLECTA, GEN. N., SP. N. FROM PERU
(NYMPHALIDAE: ITHOMIINAE )?

GERARDO LAMAS

Museo de Historia Natural “Javier Prado,” Universidad Nacional Mayor de San
Marcos, Apartado 1109, Lima-100, Pert

ABSTRACT. Paititia neglecta, gen. n., sp. n., from Peri, San Martin, Juanjui,
is described herein. This monotypic genus is considered to be the most primitive
member of the tribe Mechanitini of the Ithomiinae.

Two specimens, representing a new genus and species of Ithomiinae, lay
unnoticed for almost 50 years in the collections of the British Museum
(Natural History ), London, and the American Museum of Natural His-
tory, New York. The BMNH female was included among a series of un-
identified Methona examples in the Rothschild collection, while the
AMNH male had been identified (and labelled) as “Xanthocleis ino” by
the late R. M. Fox. The third known specimen (a male) was presented
to the “Javier Prado” Museum in 1976 by the collector, Mr. José M.
Schunke.

Paititia Lamas, new genus

External diagnostic characters. Large Mechanitini (Fox, 1956), very similar in size
and color pattern to Thyridia psidii ssp. and Methona spp. May be distinguished from
Thyridia psidii (Linnaeus) by the absence of small red dots on base of forewing above,
and the presence cf two separate, white spots on cell Sc-R:-R; of hindwing below.
Paititia may be separated from Methona spp. by the long hair patch extending beyond
the discal cell apex of the male hindwing above, which in the latter is restricted to the
basal half or two-thirds of that cell.

In Fox’s key (1940), the males of Paititia will key out to Xanthocleis
Boisduval (i.e., Thyridia Hiibner), but may be readily differentiated by
the characters given above. The females will key out to Athesis Double-
day; however, they can be easily distinguished by the wing shape and

1 The present paper forms part of a D.Sc. dissertation submitted to the Departamento de Zoologia,
Instituto de biociéncias, Universidade de S40 Paulo, Sao Paulo, Brasil (Lamas, 1973). As a con-

densed version of that thesis will not be published in the near future, I have decided to make
available the new genus and species of Ithomiinae described therein.

bo

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fics. 1-11. Paititia neglecta, gen. n., sp. n.: 1, palpus; 2, male foreleg; 3, fe-
male foreleg; 4, male forewing; 5, male hindwing; 6, female hindwing; 7, male geni-
talia, ventral view; 8, male genitalia, lateral view; 9, penis; 10, male eighth tergite, dor-
sal view; 11, female genitalia, ventral view.

color pattern (wings long and narrow, with yellowish transparent areas
in Paititia, short and wide, with reddish-brown translucent areas in
Athesis ).

Palpi (Fig. 1). Basal segment curved, adpressed to the head; second free, 1.5 times
longer than the basal; apical fusiform, very small, one-fifth to one-eighth the length of
the basal (longer in male).

Antennae. Club yellow, with approximately 15 segments, very slightly widened,

VOLUME 33, NUMBER 1 3

barely more so than the pedicel, which is black; as long as the forewing discal cell,
that is, slightly longer than one-half the forewing length.

Male forelegs (Fig. 2). Reduced; femur plus trochanter as long as the stout
articulate coxa; tibia one-fourth longer than femur; tarsus two-fifths the length of the
tibia, with slender spines on ventral side.

Female forelegs (Fig. 3). Reduced, coxa articulate; femur as long as coxa; tibia
equal to femur plus trochanter; tarsus with 4 apparent segments, two-fifths the length
of the tibia; basal segment twice the lengh of the remainder; second and third short,
apical consisting of the fourth and fifth fused; first segment with two pairs of spurs,
the outer ones slightly longer; second and third segments with a pair of spurs each;
second, third and apical segments with trichoid sensillae.

Venation (Figs. 4-6). Sc and R: on forewing of both sexes running parallel, with-
out anastomosing, Sc ending shortly beyond discal cell apex; Ri arising from cell, base
of Re anastomosed to Rs-s-s, well beyond cell apex; rs-m: absent or very short, straight;
mi-mz long, angular, long Re arising from angle; msz-ms also long, straight.

Male hindwing with bifid hum, both arms well developed; Sc and R separate at
base, Sc ending beyond discal cell apex; rs-m: straight, slightly more than half as
long as mms; mi-mz angular, both arms straight and of equal length, Rc on angle;
Me2-ms straight. Hair patch complete, running along upper portion of discal cell, behind
radial vein, from base to beyond cell apex.

Female hindwing. Similar to male, except that Sc ends scarcely beyond cell apex;
rs-mi very short, less than one-third the length of m2-ms; no hair patch.

Male genitalia (Figs. 7-10). Eighth tergite with two short and distally widened
lateral lobes, without sclerotized claws; saccus short, one-half as long as the valva;
tegumen hood-like, not separated by a suture from uncus; uncus stout, posterior area
with a strong claw bent downwards; juxta V-shaped, poorly developed; appendices
angulares not sclerotized; gnathos lightly developed, its arms being united below the
tuba analis by means of a membrane only; valvae symmetrical, long, quite wide, bear-
ing two claw-like appendices on their caudal ends, upper claw shorter and stronger
than lower; penis long, slender, curved between the anterior and middle thirds; fora-
men penis very long, one-third of the total length; gonoporus flared, terminal; vesica
with cornuti.

Female genitalia (Fig. 11). Vaginal plate irregular, funnel-shaped, with a lateral
aliform process on left side; ostium bursae located left of longitudinal axis of abdomen;
caudal end of ostium bursae strongly sclerotized and slightly bent left and downwards,
remainder of ductus very long; bursa copulatrix with poorly developed signa and with
a globular appendix bursae.

Type-species: Paititia neglecta Lamas, sp. n.

Etymology. The generic name is based on the Quechua word “‘Paititi,” which re-
fers to the fabulous kingdom of “El Dorado,” so ardently sought by explorers and ad-
venturers in South America, almost ever since the discovery of the New World. It
should be treated as being of feminine gender.

Paititia neglecta Lamas, new species

Male (Fig. 12). Wing margins and transverse bands black, transparent areas with
a yellowish tinge. Hindwing below with two white spots on cell Sc + Ri-R; and a row
of marginal double white spots on cells M:-M>2 to Cu.-2A. Humeral spot white, costal
line pale yellow. Abdomen below, and all other body spots, white.

Female (Fig. 13). Similar to male, hindwing white, marginal spots smaller.

Types. HoLorypPe @, Juanjui, San Martin, Peru; xi.34 (G. Klug), deposited in the
British Museum (Natural History). One PARATYPE ¢, Achinamiza, San Martin, Peri;
14.1.26 (H. Bassler, station F6001); AMNH Acc. 33591, in the American Museum of
Natural History, New York. One PARATYPE ¢, Iberia, Madre de Dios, Peri; 27.vi.75
(J. M. Schunke ), in the Museo “Javier Prado,” Lima.

Etymology. The specific name is the Latin word for “forgotten.”

4 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fics. 12-13. Paititia neglecta, gen. n., sp. n.: 12, paratype male, Achinamiza;
13, holotype female. Dorsal left, ventral right.

DiIscussION
Relationships

Paititia is considered to be the most primitive member of the tribe
Mechanitini (Lamas, 1973). It indeed seems to represent a link between
the Tithoreini and Mechanitini.

Paititia neglecta resembles members of the genus Olyras Doubleday (cf.
Fox, 1940, 1956), in the shape of the male forelegs, wing venation and
genitalia, and even the presence of the white spots on the hindwing cell
Sc + Ry-R, below (present in most forms of Olyras). However, Olyras
differs by the divided hair patch and by the valvae having only one short
claw-like process, the females being even more different.

Paititia is very close to Thyridia Hiibner (= Xanthocleis Boisduval),
both having almost identical venation, but I believe the differences pre-
sented by the male and female genitalia and foretarsi are enough to sep-

arate them.

VoLUME 33, NUMBER 1 5

Natural history

Nothing is known about the life-habits of Paititia neglecta. Two of the
localities where it has been found (Juanjui and Iberia) are characterized
by rather dry tropical forest. This may represent an example of a primi-
tive species which has been displaced to marginal habitats by a more
modern and aggressive species, as seems to be the case with Melinaea
mnasias ( Brown, 1977).

ACKNOWLEDGMENTS

Research for the present paper was mainly financed by grants “Bio-
légicas 70/671” and “72/849” of the Fundacéo de Amparo a Pesquisa do
Estado de Sao Paulo (FAPESP), Brasil. I acknowledge with special
thanks the help and hospitality of Mr. R. I. Vane-Wright (British Museum
[Natural History] ), Dr. F. H. Rindge (American Museum of Natural His-
tory, New York), and the advice of Drs. P. E. Vanzolini and N. Papavero
(Museu de Zoologia da USP, Sao Paulo) and K. S. Brown, Jr. ( Universi-
dade Estadual de Campinas, Sao Paulo). I am deeply obliged to Mr. J. M.
Schunke for having obtained the third known specimen of Paititia neglecta
for the “Javier Prado” collections.

LITERATURE CITED

Brown, K. S., Jr. 1977. Geographical patterns of evolution in Neotropical Lepi-
doptera: differentiation of the species of Melinaea and Mechanitis (Nymphali-
dae, Ithomiinae). System. Entomol. 2: 161-197.

Fox, R. M. 1940. A generic review of the Ithomiinae (Lepidoptera: Nymphali-
dae). Trans. Amer. Entomol. Soc. 66: 161-207.

1956. A monograph of the Ithomiidae (Lepidoptera). Part I. Bull. Amer.

Mus. Nat. Hist. 111: 1-76.

1967. A monograph of the Ithomiidae (Lepidoptera). Part III. The tribe
mechanitini Fox. Mem. Amer. Entomol. Soc. 22: 1-190.

Lamas, G. 1973. Taxonomia e evolucdo dos géneros Ituna Doubleday (Danainae) e
Paititia, gen. n., Thyridia Hiibner e Methona Doubleday (Ithomiinae) ( Lepidop-
tera, Nymphalidae). D.Sc. Thesis to the Departamento de Zoologia, Instituto de
Biociéncias, Universidade de Sao Paulo, vii + 225 p.

Journal of the Lepidopterists’ Society
33(1), 1979, 6-20
EXPERIMENTAL HYBRIDIZATION BETWEEN PHYCIODES
THAROS AND P. BATESII (NYMPHALIDAE )

CHarLES G. OLIVER!
R. D. 1, Box 78, Scottdale, Pennsylvania 15683

ABSTRACT. F: hybrids and backcrosses were made between the nymphalid but-
terflies Phyciodes tharos and P. batesii. The two species differ in larval, pupal, and
adult phenotypic appearance, ecology, and larval diapause response. Genetic in-
compatibility was shown by significant hybrid inviability, growth irregularities, and
abnormal adult sex ratios and development times. The reciprocal F: hybrids differed
greatly in their expression of incompatibility. Hybrid inviability is attributed to break-
downs in the genetic mechanisms controlling growth and development.

The relationship of Phyciodes tharos Drury to P. batesii Reakirt has
been little understood despite the fact that they both occur in compara-
tively densely populated areas of the northeastern United States and
have been known to be distinct for well over a century. This confusion
appears to be due to two causes. First, P. tharos is common to abundant
over the entire range of P. batesii. Since P. batesti occurs in widely sepa-
rated, small populations, it is probably often overlooked due to its super-
ficial resemblance to P. tharos. Second, the biology of P. tharos itself actu-
ally is poorly known. Rearing and hybridization studies now in progress in
my laboratory indicate that “P. tharos” in the Northeast consists of two
entities differing in larval and adult phenotypic appearance and voltinism
and showing significant incompatibility when hybridized in the laboratory
(Oliver, in prep.). The more southern entity, hereafter referred to as
“Type A,” occupies the Transition and Austral Life Zones, whereas the
more northern “Type B” is the “P. tharos” of northern New England,
northern New York State, and southern Canada. The phenotypic differ-
ences of Types A and B have resulted in Type B passing as P. batesii in
many collections, although it resembles P. batesii little more than does
Type A.

Phyciodes batesti is very local in the Northeast. It appears to be restricted
to dry sites, often of the barrens type. The Onondaga Co., New York, pop-
ulation used in these experiments occurs on dry limestone ledges. In —
the Appalachians of Pennsylvania and West Virginia, the localities I
have investigated are for the most part shale barrens or rocky riparian
slopes. One of the best known localities, along the banks of the Ottawa
River near Aylmer, Quebec, was described by McDunnough (1920)
as the “lower dry slopes of a small ridge.” Both Types A and B of P. tharos
occur in a wide variety of habitats, including those of P. batesii.

7 Adjunct. Assistant Professor, Dept. of Biology, West Virginia University, Morgantown, West
Virginia 26506.

VoLUME 33, NUMBER 1 t

Phyciodes batesii flies in early June in West Virginia and southwestern
Pennsylvania and in mid June in central New York State. In West Virginia
and southern Pennsylvania this is between the rather discrete first and sec-
ond broods of P. tharos Type A, which has a total of three to four broods.
In Onondaga Co., New York, however, P. batesii and P. tharos Type B fly
together. Thus, the observation of Forbes (1960) that broods of P. tharos
and P. batesii alternate at any given locality seems to apply only to the
southern portion of the range of P. batesii.

The life history of P. batesii was described and figured by McDun-
nough (1920). The larva and pupa differ in a number of characters from
those of P. tharos (Table 1) and are much more like those of P. campes-
tris, which were figured by Comstock (1930) and compared briefly to
P. tharos in another paper of mine (Oliver, 1978). First and second instar
larvae of both P. campestris and P. batesii live communally within a loose
web spun over the feeding area on the foodplant. P. tharos has a similar
communal behavior, but no web is spun.

McDunnough was able to obtain oviposition by P. batesii on “a species
of Aster with heart-shaped leaves,” found wild larvae on this plant, and
successfully reared them through to adults. This Aster was probably A.
undulatus L., which is very common in P. batesii habitats. Wild-caught
P. batesii females from Onondaga Co., New York, refused to oviposit on
Aster undulatus in the laboratory, but laid readily on A. simplex ( Willd.)
Burgess, a common foodplant of P. tharos. Newly-hatched larvae of both
Phyciodes fed readily on A. undulatus when transferred to its leaves.
Larvae of both species also accepted A. laevis L.

More than 200 unmated, laboratory-reared P. batesii adults were re-
leased shortly after eclosion during August into a western Pennsylvania
old-field habitat containing abundant Aster simplex (but no A. undula-
tus). Several pairs were observed in copulo the next day. In early Octo-
ber a group of small larvae was recovered from a clump of A. simplex and
reared through to normal adults the following spring, indicating that P.
batesii may choose more than one species of Aster as natural foodplants.

In the northern Midwest (e.g., Michigan and Wisconsin) P. batesii has
a somewhat different biology. Colonies occur in moist areas, and there is
sometimes a partial second brood (Nielsen, in litt.). Populations in the
Northeast seem to be strictly univoltine. Midwestern P. batesii differ
slightly in appearance from those in the Northeast and may possibly rep-
resent a separate entity.

PROCEDURE

Stock of P. batesii used in these experiments was derived from four
wild-inseminated females collected 11 June 1976, in Syracuse, Onondaga

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

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-VoLUME 33, NUMBER 1 9

Co., N.Y. Stock of P. tharos Type A was derived from four females taken
10 June 1976, in Acme, Westmoreland Co., Pennsylvania, and of Type B
from two females taken 11 June 1976, at the Onondaga Co., N.Y. locality.
Cultures were maintained at 15 to 28°C and separated into controlled
photoperiod (18 h light/24 h or 24 h light/24 h) and natural photoperiod
(for Fayette Co., Pa., latitude 40°N) groups. Artificial lighting after
sunset was provided by a 100-watt incandescent bulb at a distance of 1
to 2 meters.

Matings were made by the hand-pairing method (Clarke, 1952).
Fecundated females were provided with cut sprigs of Aster simplex for
oviposition. Eggs were left in situ until hatching and the aster sprigs kept
fresh in water. Larvae were reared on cut sprigs of A. simplex in water
and housed in 10 X 20 cm glass cylinders.

The photoperiodically-regulated larval diapause in Phyciodes occurs at
the beginning of the third instar. Diapausing larvae were removed from
active cultures, placed in groups in 90 mm plastic petri dishes, sealed in
airtight containers, and stored in a domestic refrigerator at 0 to 2°C until
April or May of the following year. Upon removal from the diapause con-
tainers al] larvae were maintained at 15 to 28°C and 18 h light/24 h until
pupation, at which time they were transferred to natural light conditions.

F, progeny of wild-collected females was used for the hybrid pairings
and as parental-type stock for backcrosses; no stock used was inbred. Ob-
servations were made on parental population and F,; hybrid phenotypic
appearance, interspecific courtship behavior, development periods and
adult eclosion patterns, fertility, adult sex ratios, embryonic, pupal, and
adult viability, and on backcross embryonic viability. Controls were
reared concurrently at all times for comparison with experimental broods.

Data on egg fertility, viability, and sex ratios were treated statistically
using the Wilcoxon Two-sample Test. Adult fertility was measured by a
count of the number of visibly developing eggs divided by the total num-
ber of eggs laid after a single mating. Development periods from hatch-
ing of the egg to eclosion of the adult were estimated by calculating the
99% confidence intervals for the medians of the distributions (Owen,
1962). Distributions of development times within broods or series of
broods were represented by adult eclosion curves, graphs of the number
of adults eclosing from pupae each day.

RESULTS

Interspecific courtship behavior

Courtship in both P. tharos and P. batesii is apparently dependent on a
variety of stimuli. The presence of a butterfly of appropriate coloration

10

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

VoLUME 33, NUMBER 1 Wl

and size elicits approach by males. If the approached individual does not
leave or, in the case of conspecific males, show aggressive behavior, as
attempt at copulation will be made. Females which do not wish to cop-
ulate avoid the male’s probing genitalia by dorsal or lateral movement of
the abdomen. Males of both P. tharos and P. batesii showed the courtship
approach response when presented with females of the other species. After
an initial response, however, males of neither species attempted to copu-
late. Some stimulus, perhaps olfactory, appeared to terminate courtship.

Phenotypic appearance

Differences in phenotypic appearance of the fifth instar larvae, pupae,
and adults of the parental species and F, hybrids are summarized in Table
1. Typical specimens of adults are shown in Fig. 1. There was wide varia-
tion in adult phenotypic appearance among the F, hybrid broods.

The artificial illumination levels used to extend daily photophase in
these experiments eliminated facultative diapause in P. tharos (see
below ), but the induction of the naturally-occurring adult seasonal forms
was little affected. The photoperiodic regulation of polyphenism in P.
tharos has been described in a previous paper (Oliver, 1976). Univoltine
P. batesii do not, of course, show natural seasonal polyphenism. Under the
artificial long-day laboratory conditions described above, however, non-
diapausing larvae produced late summer and early fall adults which had
significantly heavier expression of the dark wing pattern elements on both
the ventral and dorsal sides. This artificially induced form did not differ
from the naturally occurring phenotype as the seasonal forms of P. tharos
differ from each other. F, hybrid adults emerging in September and Octo-
ber showed some expression of the short photophase phenotype but not to
as great an extent as did the P. tharos controls.

Phyciodes tharos and P. batesii differ in the length of time required for
full embryonic development. Eggs of P. tharos kept at natural outdoor
temperatures during early August hatched after 6 days. Those of P. batesii
required an additional day or day and a half. The hatching times of the F,
hybrids varied within broods from 6 to 7% days.

Fertility, viability, and sex ratio

Egg fertility and embryonic viability of control broods was very high
(Table 2). Fertilizability of eggs was slightly reduced in the F, hybrid

<<

Fic. 1. Parental-type and F, hybrid adults: Row A—P. tharos; B—P. batesii;
C—F;, hybrids P. batesii 2 x P. tharos 6; D—F: hybrids P. tharos 2 x P. batesii ¢.
Specimens show, left to right, male dorsal, female dorsal, male ventral, female ventral.

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

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VOLUME 33, NUMBER | 13

crosses. Embryonic viability was heavily depressed in the P. batesii 2 x
P. tharos 6 F, hybrid series, but not demonstrably affected in the recip-
rocal cross. Embryonic viability of the P. batesii 2 x N.Y. P. tharos é Fy
hybrid series may differ from that of the P. batesii 2 X Pa. P. tharos ¢
series (P = .10). Backcross embryonic viability was drastically reduced.
Embryonic viability of both F; hybrids and backcrosses showed great
variability among broods (Table 2).

Post-larval viability (i.e. during pupation and eclosion and as pupae)
was greatly reduced in the P. batesii 2 x P. tharos é F, hybrids and to a
lesser extent in the reciprocal cross. The decrease in viability was almost
entirely during the pupal stage (Table 3).

Adult sex ratios of the P. batesii 2 x P. tharos 6 F, hybrid series did not
differ significantly from those of the parental broods (Table 3). In the
two P. tharos 2 x P. batesii é broods, however, female adults were en-
tirely absent (Brood 76-43) or greatly reduced in numbers (to 16.07% in
Brood 76-42).

Structural abnormalities

Structural abnormalities involving segmental irregularities were rela-
tively common in the P. batesii 2 x P. tharos 6 hybrid broods, though
absent in the parental controls and reciprocal crosses. Between 0 and
15% (no exact counts made) of the larvae of each hybrid brood showed
a lack of development of one side of an abdominal segment. The af-
fected segment half was both narrower and shorter than the correspond-
ing half, often lacked a tubercle, and resulted in the larval abdomen
abruptly bending to one side. Two larvae were segregated and observed
throughout development. In these, the semental irregularity persisted
through the pupa and into the adult (Fig. 2). Both larvae produced ap-
parently otherwise normal male adults.

F, hybrid males from the cross P. batesii 2 <x P. tharos é had dispropor-
tionately large, flaccid abdomens (Fig. 1). The genitalia of many in-
dividuals were apparently permanently extruded, and hand-pairings us-
ing any males were very difficult. No males would mate naturally in
cages.

Voltinism

The P. tharos Type A population culture from Westmoreland Co., Pa.,
showed no incidence of larval diapause when reared on a photoperiod re-
gime of 18 h light/24 h or under natural photoperiod conditions during
June and July. Broods reared under natural photoperiod during August
and September showed a significant incidence of diapause (Table 4).
The P. tharos Type B population culture from Onondaga Co., N.Y., on the

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

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VoLuME 33, NUMBER 1

Fic. 2. P. batesii 2 x P. tharos 6 F, hybrid male adult showing asymmetrical
development of abdominal segment (arrow).

16 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TABLE 5. Development times in days from hatching of egg until eclosion of adult
for non-diapausing P. tharos and P. batesii control broods, Fi hybrids, and backcrosses.
Medians with 99% confidence. See text for rearing conditions.

Brood Date Males Females
no. hatched N Min—Max Median N Min—Max Median

P. tharos (New York )—Natural photophase

76-8 19-26 June WH 32-53 34-36 134 33-52 38-39
76-9 19-23 Jun 216 32-50 34 190 32-41 36-37
P. tharos (New York )—18 h light/24 h
76-38 13 Aug 84 33—44 36-37 95 37-54 42-44
76-40 13 Aug 90 33-46 34-35 79 35-70 37-42
76410 | 13 Aue Bee 38/1 31 | teams =39
P. batesii—24 h light/24 h
76-1 21-25 Jun Di 38-107 49-56 on 43-116 56-57
76-2 Bilis) |own 54 33-105 49-76 49 40-116 55-76
76-3 24-25 Jun 23 40-97 48-55 34 48-134 51-62
76-4 21-25 Jun 30 38-86 41-64 33 38-89 45-55
P. batesii 2 x P. tharos 6—18 h light/24 h
76-5 13-15 Aug 11 40-61 42-59 8 35-54 35-54
76-6 14-18 Aug 61 39-73 49-53 65 33-56 41-46
76-7 15-18 Aug 15 ATT 51-67 D4 34-65 42-47
76-8 20-21 Aug 4 Silas} 51-58 5 39-56 39-56
76-10 15-21 Aug 40 45-74 51-54 on 35-55 39-42
foal 22-29 Aug 28 44-65 49-57 19 38-57 41-48
76-13 22-23 Aug 3 59-69 — iL 50 —
(Fl 4 Jun 4 44-84 = 9 3440 34—40
77-2 30 May-2 Jun 24 40-60 42—A7 18 36-48 36—40
77-3 3-5 Jun 10 34-41 34-41 6 32-36 —
P. tharos 2 x P. batesii 6 —18 h light/24 h

77-42 15-18 Aug 188 35-62 41-42
77-43 16-17 Aug AO 36-58 38-40
(P. tharos 2 x P. batesii 6)? x P. tharos 6—18h light/24 h
77-26 19-20 Jul 42, 28—46 29-34 54 29-45 37-38
77-34 28-29 Jul 110 30-54 34-37 97 29-37 30-31
77-38 31 Jul-l Aug 79 29-41 33-35 106 32-44 36-37
P. tharos 9 x (P. batesii 2 x P. tharos 4 ) 6 —18 h light/24 h
7728) 29-31) Tul 18 30-645 e0eso

77-32 24-27 Jul 164 27-66 33-35 97 28-65 31-36

other hand, showed significant incidence of larval diapause when reared
during June and July on natural photoperiod. This diapause response was
entirely facultative, since there was a complete absence of diapause in
larvae reared on 18 h light/24 h.

Phyciodes batesii reared on natural photoperiod during June and July
showed a 100% incidence of larval diapause. Many of those reared on 24 h

VoLUME 33, NUMBER 1 17

NUMBER OF ADULTS ECLOSING

SES yp
116 134

DAYS UNTIL ECLOSION

Fic. 3. Distributions of times required for development of New York State P.
tharos, P. batesii, and Fi hybrid broods from hatching of eggs until eclosion of adults.
A—P. tharos hatching in late June, natural photophase, B—P. batesii hatching in late
June, 24 h light/24 h; C—F, hybrid P. batesii 2 x P. tharos é hatching in early
August, 18 h light/24 h; D—F: hybrid P. tharos 2 x P. batesii & hatching in early
August, 18 h light/24 h.

light/24 h, however, developed without diapause, indicating that at least
part of the culture was composed of facultatively diapausing individuals.

There was no incidence of diapause in F, hybrid larvae from the cross
P. batesii 2 X P. tharos ¢. The reciprocal hybrid, however, had an inci-
dence of diapause intermediate between those of the parental species
(Table 4). Survival of the F; hybrid larvae during diapause storage was
normal compared with that of the parental species.

Development periods and eclosion patterns

The median development periods from hatching of the egg to eclosion
of the adult were significantly longer for non-diapausing P. batésii than
for concurrently reared non-diapausing P. tharos from New York State or
Pennsylvania (Table 5). In addition, emergences were very scattered,
producing a very different eclosion pattern from that of either population
of P. tharos ( Figs. 3A and B).

18 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

TaBLE 6. Post-diapause development periods of P. tharos, P. batesii, and their Fi
hybrid P. tharos 2 x P. batesii 6. Medians with 99% confidence. All larvae re-
moved from storage on same day (4 April 1977).

Brood Males Females

no. N Min—Max Median N Min—Max Median
P. tharos (New York)
76-8 105 41-49 43-44 153 42-56 49-50
P. batesii

76-1 1 46 — 8 44-50 44-50
76-2 5 46-48 — i 49-51 49-51
76-3 14 43-A7 43-46 13 45-54 45-48
76-4 4 45-51 — 5 48-53 —

P. tharos @ x P. batesii 3
76-42 = 36 72—107 74-83

F, hybrids of the cross P. batesii 2 X P. tharos é showed eclosion pat-
terns intermediate between P. tharos and P. batesii, but male develop-
ment periods about the same as those of P. batesii. Females of this cross
had shorter development periods and tended to emerge before the males,
rather than the normal reverse (Fig. 3C).

Male development times of the cross P. tharos ? X P. batesii 6 were
significantly shorter, though not as short as those of concurrently reared
P. tharos. The spread of eclosion times for male adults was much more
like that of P. tharos than P. batesii (Fig. 3D). All P. tharos ? X P. batesii
6 F, adults emerging without diapause were males; all those emerging
after diapause, females. Resumption of feeding after diapause of these
larvae was much delayed after removal from cold storage, and growth
was much slower than that of the parental controls (Table 6).

DIscussION

It is clear from the results that P. tharos and P. batesii are well-differ-
entiated species. There are marked differences in phenotypic appearance,
voltinism, development rate, and ecology. The adults show strong be-
havioral isolating mechanisms during courtship in the laboratory. The
heavy reduction in F, hybrid viability and fertility and in backcross em-
bryonic viability indicates a high degree of genetic incompatibility be-
tween the species.

Phyciodes tharos and P. batesii appear to have achieved rather similar
phenotypes (compared to P. tharos and P. campestris, for example) by
somewhat different genetic means. This is attested to by the wide range of
phenotypic variation in the F; hybrid adults and is especially marked in
F, females from the cross P. batesii 2? x P. tharos 6. Expression of the

VOLUME 33, NUMBER 1 19

dorsal dark pattern elements ranges from as dark or darker than P. batesii
to almost as light as P. tharos (Fig. 1). This indicates that the two species
look more alike than they actually are.

I have discussed at length in another paper (Oliver, in press) the
genetic basis of incompatibility effects involved in hybrid breakdown and
surveyed the literature on viability of butterfly hybrids. The hybrid in-
compatibility shown between P. tharos and P. batesii may involve differ-
ences in the genetic control of hormones that direct growth and develop-
ment. Disruption of normal hormonal control leads to formation of
inviable embryos, abnormal tissue differentiation patterns, lessened fer-
tility, lowered ability to pass from one life cycle stage to another, and ab-
normal development rates.

In general the genetic incompatibility between P. tharos and P. batesii
is fairly similar to that shown between P. tharos and P. campestris mon-
tana Behr (Oliver, 1978). Both sets of F; hybrids have similar adult
eclosion patterns. However, embryonic viability is reduced much more
in the series P. tharos x P. batesii than in P. tharos X P. campestris. On
the other hand, there is a much greater deficiency of F; hybrid females in
the series P. tharos X P. campestris than in P. tharos x P. batesii. These
differences and those in ecology, voltinism, and so on indicate that P.
batesii and P. c. montana are physiologically quite distinct, and P. batesii
probably should not be regarded as an eastern representative of P. cam-
pestris. P. batesii is more specialized than either of the other species. It
appears to have evolved from multivoltine stock by lowering of the thresh-
old of diapause induction to include all naturally encountered photoperiod
conditions. Although capable of feeding on at least several species of
Aster, it has become closely associated with (though perhaps not re-
stricted to) a single species, A. undulatus, and seems in the Northeast to
be found only in the rather narrow habitat range of this aster. Other but-
terflies have followed similar courses of evolution. Pieris virginiensis
Edw., for example, has evolved univoltinism in apparently the same way
(Shapiro, 1971; author's unpub. data) and has become restricted to the
narrow habitat of its single foodplant, Dentaria diphylla Michx.

ACKNOWLEDGMENTS

I greatly appreciate the help of Mr. Edward Jennejohn, Manlius, New
York, in obtaining laboratory stock of Phyciodes batesii. Mr. Joseph O.
Brenneman provided invaluable photographic assistance.

LITERATURE CITED

Ciarke, C. A. 1952. Hand pairing of Papilio machaon in February. Entomol. Rec.
64: 98-100.

20 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

Comstock, J. A. 1930. Egg, larva, and pupa of Phyciodes campestris Behr. Bull.
So. Calif. Acad. Sci. 29: 136.

Forses, W.T.M. 1960. Lepidoptera of New York and neighboring states. Part IV.
Memoir 371, Cornell U. Agric. Exper. Sta.

McDunnovucp, J. 1920. Notes on the life history of Phyciodes batesi Reak. (Lepid. ).
Canad. Entomol. 52: 56-59.

Outver, C.G. 1976. Photoperiodic regulation of seasonal polyphenism in Phyciodes
tharos (Nymphalidae). J. Lepid. Soc. 30: 260-263.

1978. Experimental hybridization between the nymphalid butterflies

Phyciodes tharos and P. campestris montana. Evolution 32: 594-601.

1979. Genetic differentiation and hybrid viability within and between
some Lepidoptera species. Am. Nat. (in press ).

Owen, D. B. 1962. Handbook of statistical tables. Addison-Wesley Publ. Co.,
Reading, Mass.

SHAprro, A. M. 1971. Occurrence of a latent polyphenism in Pieris virginiensis
(Lepidoptera: Pieridae). Entomol. News 82: 13-16.

Journal of the Lepidopterists’ Society
33(1), 1979, 20

TEMPORARY RANGE EXTENSION AND LARVAL FOODPLANT OF
DYNAMINE DYONIS (NYMPHALIDAE) IN TEXAS

The northern normal limit of Dynamine dyonis Geyer in Texas is Cameron and
Hidalgo counties. The occurrence of D. dyonis north of its usual range was noted on
27 July 1966, when I collected a tattered female on the Salado Creek, three miles
southeast of the Northeast Preserve, a city park in San Antonio, Texas. The inunda-
tions of Hurricane Beulah in September 1967 produced lush vegetation in south Texas
and may have caused the invasion of D. dyonis to extend as far north as Collin Co.
(18 Sept. 1968, 2 ¢ 6, leg. Edward Reid). Further evidence of this movement are
records by the following collectors in other counties in 1968: Gonzales Co., Hidalgo
Co. (both M. A. Rickard); Bell Co., San Patricio Co. (both R. O. and C. A. Kendall);
Travis Co. (C. J. Durden); Brazos Co. (J. E. Hafernik). The last known record in
1968 was Bexar Co., 23 Nov., 1 @, leg. J. F. Doyle. The total number of D. dyonis
taken by collectors in Texas from 27 July 1966 through 23 Nov. 1968 was 134 (68
64, 66 292). To my knowledge no populations remain in central or northern
Texas.

On 5 May 1968, I observed a female D. dyonis as it fluttered about a trailing plant
in a dry creek bed in the Northeast Preserve. The plant, Tragia ramosa Torrey (Eu-
phorbiaceae), and the butterfly were caged and placed outdoors at my home in San
Antonio, Texas. Twenty-four eggs were deposited that same day. The first larva
emerged on 9 May. Only 6 larvae remained on 19 May because of cannibalism. Adults —
which emerged were: (31 May) 3 6 6,1 9; (1 Jume) 1 6,1 @.

Larvae were collected at the Northeast Preserve site in 1968 and reared on T.
ramosa. These larvae were collected on 19 May and pupated 24 May. One adult ( é )
emerged on 5 June. Larvae also were collected on 14 July and pupated between 17
and 18 July. Adults (1 6, 1 2) emerged on 23 and 25 July.

Josepn F, Doyxe III, 11839 Monticeto Lane, Stafford, Texas 77477.

Journal of the Lepidopterists’ Society
33(1), 1979, 21-28

NOMENCLATORIAL CHANGES IN EUCOSMINI
(TORTRICIDAE)

RicHARD L. BROWN
Department of Entomology, Comell University, Ithaca, N.Y. 14853

ABSTRACT. The genera Kundrya, Norma, and Erinaea are synonymized with
Rhopobota. Female genitalia of R. unipunctana, R. dietziana, and R. finitimana are
illustrated. Griselda stagnana and G. myrtillana are transferred to Rhopobota. Epi-
blema separationis, formerly a subspecies of E. praesumptiosa, is recognized as a
species. Female genitalia of both species are illustrated. Notocelia trimaculana, N.
illotana, N. culminana, and N. purpurissatana are distinguished from Epiblema.

Problems of identification, classification, and evolutionary relationships
of the Olethreutinae have persisted in spite of the economic importance
of many species in this subfamily. Heinrich’s revision (1923) of the
Eucosmini was based principally upon characteristics of wing venation
and male genitalia. In recent years Bentinck and Diakonoff (1968),
Diakonoff (1973), and Obraztsov (1958-1968), have recognized the value
of the female genitalia in differentiating species as well as defining the
genera. This paper presents modifications in the classification of selected
Eucosmini genera and species as a result of the examination of the female
genitalia and other characters.

Changes in Rhopobota

Rhopobota Lederer, 1859, Wien. Ent. Monat., 3: 366.

Type species: Tortrix naevana Huebner [1814-1817], by monotypy. Although
Lederer considered R. naevana a senior synonym of Tortrix unipunctana Haworth
[1811], R. naevana is now recognized as a junior subjective synonym of R. unipunc-
tana.

Norma Heinrich, 1923, U.S. Natl. Mus. Bull. 123: 191. [New Synonymy. ]

Type species: Epinotia dietziana Kearfott, 1907, by monotypy.

Kundrya Heinrich, 1923, U.S. Natl. Mus. Bull. 123: 192. [New Synonymy.]

Type species: Kundrya finitimana Heinrich, 1923, by monotypy.

Erinaea Meyrick, 1907, Journ. Bombay Nat. Hist. Soc., 18: 141. [New Synonymy. ]

Type species: Erinaea chlorantha Meyrick, 1907, by monotypy; a junior subjec-
tive synonym of Teras verditer Hampson, 1891 (Diakonoff, 1950).

The genera Norma and Kundrya were considered by Heinrich to be
close to Rhopobota. Heinrich distinguished Kundrya by a character of
the forewing venation, Ry and R; united. These two veins were described
as stalked in Rhopobota and approximate in Norma. Rhopobota was
separated from the first two on the basis of the porrect socii which are

22 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Zug

en gen ae
ea

2

Fics. 1-2. Female genitalia including seventh abdominal segment: (1, left)
Rhopobota unipunctana (Bellingham, Washington; USNM 17763); (2, right) Rhopo-
bota dietziana (Ethel, Arkansas; R. L. Brown prep. 683). Scale line = 1 mm.

apically fused. Meyrick (1907) described Erinaea based upon a species
from Ceylon, and did not indicate a relationship with Rhopobota.
Unique characters of the female genitalia of all the type species and a
reassessment of previous distinctions provide the basis for synonymizing
Erinaea, Norma, and Kundrya with Rhopobota. The most distinctive
character is the sclerotization of the sides and base of the corpus bursae
and distal area of the ductus bursae. A separate sclerotized band around
the ductus bursae is located near the colliculum at the inception of the

VoLuME 33, NUMBER 1 93

Fic. 3. Female genitalia including seventh abdominal segment of Rhopobota
finitimana (Falls Church, Virginia; USNM 17762). Scale line = 1 mm.

ductus seminalis. The colliculum is sclerotized ventrally and extends be-
yond the lamella antevaginalis. Both the lamella antevaginalis and lamella
postvaginalis are well developed and may be fused with or separate from
the seventh sternite (Figs. 1, 2, 3; Clarke, 1958, pl. 169). These charac-
teristics are also shared by the Asian species, R. eclipticodes (Meyrick)
and R. microrrhyncha (Meyrick), as figured by Clarke (1958, pl. 170)
and the Palearctic species, R. ustomaculana (Curtis), figured by Bentinck
and Diakonoff (1968, fig. 189b, c).

The eighth tergite which surrounds the papillae anales possesses scales
as well as simple setae in the three North American species, dietziana,
finitimana, and unipunctana. Scales may be present or absent on the
eighth tergite among the species of the related genera, Epinotia, Ancylis,
and Chimoptesis.

24 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

The male genitalia of finitimana, dietziana, and unipunctana are simi-
lar in having a bifurcate uncus with widely separated arms that are fin-
gerlike and weakly sclerotized. The socii are long, porrect, and densely
setose, separate in finitimana and dietziana and fused apically in uni-
punctana. The gnathos of Rhopobota is reduced and weakly sclerotized
medially. The aedeagus is short in finitimana, longer in dietziana and uni-
punctana. The male of chlorantha has not been examined.

The condition of veins Ry and R; in the forewing represents a transi-
tion from approximate in dietziana (n=17) to stalked in unipunctana
(n = 37) and united in finitimana (n =7). The length of the stalk varies
among the specimens of unipunctana examined. These veins are stalked
in the type of chlorantha, as illustrated by Clarke (1958).

All three North American species of Rhopobota feed on Ilex ( Aquifolia-
ceae). R. unipunctana also feeds on Vaccinium (Ericaceae) and is a pest
of cranberry, Vaccinium macrocarpon Aiton (Heinrich, 1923). The lar-
vae of unipunctana and dietziana were described by MacKay (1959) and
were considered to be closely related.

Analysis of photographs and drawings of the genitalia (Bentinck and
Diakonoff, 1968; Pierce and Metcalfe, 1922) provides evidence for syn-
onymizing Griselda stagnana (Denis and Schiffermueller) and G. myr-
tillana (Westwood) with Rhopobota. Obraztsov (1945) included these
two species in Griseida Heinrich, based on the presence of a costal fold
in the forewing of the male. Powell (1964) provisionally retained these
two in Griselda but indicated that they might not be congeneric with the
type species, G. radicana (Walsingham), the sole North American spe-
cies. The male and female genitalia of G. radicana are similar to those
of Epinotia hopkinsonana (Kearfott) and E. subviridis Heinrich, all coni-
fer feeders.

The presence of the costal fold in stagnana and myrtillana does not
justify their separation from the other Rhopobota species which lack one.
Secondary sexual characteristics, such as the costal fold of the male, are
seldom of generic value, as is emphasized by Diakonoff (1973) in his
study of the South Asiatic Olethreutini. My investigations of Epinotia
also show the costal fold may be present or absent in closely related spe-
cies.

Rhopobota stagnana ({Denis and Schiffermueller]). [New Combination.]

Tortrix stagnana [Denis and Schiffermueller], 1775, Ankundung eines Systemati-
schen Werkes von den Schmetterlingen der Wienergegend, p. 131.

Tortrix fractifasciana Haworth, [1811], Lepidoptera Britannica, 3: 466.

anaes fractifasciana, Pierce and Metcalfe, 1922, Genitalia British Tortricidae,
p. 75, pl. 26.

Griselda fractifasciana, Obraztsov, 1945, Zeitschr. Wiener Entomol. Ges., 30: 33-34.

The female of R. stagnana has a corpus bursae with sclerotized sides.

VOLUME 33, NUMBER 1 25

The sterigma is similar in shape to that of R. dietziana, both lamella post-
vaginalis and antevaginalis are well developed. A sclerotized band
around the ductus bursae is located near the colliculum. Scales are pres-
ent on the eighth tergite, as shown in the specimen figured by Bentinck
and Diakonoff (1968).

R. stagnana occurs in England and central Europe (Meyrick, 1895),
and feeds on the flowers and seeds of Scabiosa columbaria L. (Dipsa-
ceae ) in England ( Ford, 1949).

Rhopobota myrtillana (Westwood). [New Combination. ]

Sericoris myrtillana Westwood, In Humphreys and Westwood, 1845, British Moths,
2: 146, pl. 89, fig. 15.

Grapholitha vacciniana Zeller, 1846, Isis von Oken, p. 248.

Rhopobota vacciniana, Pierce and Metcalfe, 1922, Genitalia British Tortricidae, p.
fospl. 26.

Griselda vacciniana, Obraztsov, 1945, Zeitschr. Wiener Entomol. Ges. 30: 34.

Griselda myrtillana, Bradley, 1959, Entomol. Gazette, 10: 72, pl. 11.

The female has a similar, although weaker, sclerotization of the corpus
bursae than that described above. The sclerotized band around the
ductus bursae is located near the colliculum. Both the lamella antevag-
inalis and postvaginalis appear well developed and separate from the
seventh sternite. Scales on the eighth tergite probably are present, but
are not evident in the figures. The male has a rudimentary clasper on the
valva, similar to R. wnipunctana and ustomaculana.

R. myrtillana occurs through north and central Europe and the British
Isles. The larvae feed on Vaccinium myrtillana L. (Ericaceae) (Mey-
rick, 1895).

Changes in Epiblema Huebner

Epiblema separationis was described by Heinrich (1923) as a sub-
species of praesumptiosa Heinrich but is raised to the species level in
this paper. Heinrich characterized separationis by its smaller size, the
absence of brown spots on the inner margin of the forewing ocellus and
vein 1A, and the more rounded cucullus. E. separationis has a forewing
expanse of 9-11 mm (n=10); praesumptiosa has a forewing expanse of
14-17 mm (n=11). However, the maculation of the forewing varies
with the brown spots present or absent in each.

The most conspicuous differences are found in the female genitalia
and seventh abdominal segment (Figs. 4, 5). E. separationis lacks signa
on the corpus bursae, whereas praesumptiosa has two well developed
signa. The papillae anales of separationis are widened apically but those
of praesumptiosa are nearly uniformly wide. The lamella postvaginalis
is more setose and emarginate posteriorly in separationis. The sclerotiza-

6 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

Fics. 4-5. Female genitalia including seventh abdominal segment: (4, left) Epi-
blema separationis (San Benito, Texas; USNM 17735); (5, right) Epibelma prae-
sumptiosa (Brownsville, Texas; USNM 17708). Scale line = 1 mm.

tion of sternite VII is relatively narrower and longer in separationis than
in praesumptiosa. Both species lack sclerotization of the ductus bursae;
however, the ductus bursae of praesumptiosa is distinct in being striate.
Both species possess setae but lack scales on tergite VIII.

E. separationis possesses a tri-lobed eversible pouch ventrally, anterior
to the papillae anales. This pouch can be everted by forcing alcohol with
a syringe into the anterior opening of the seventh segment after it has
been separated from the rest of the abdomen. The pouch appears to be
glandular under high magnification.

VOLUME 33, NUMBER 1 27

The male genitalia of separationis differ from those of praesumptiosa
in having shorter setae on the corona of the rounded cucullus, a smaller
rudimentary clasper, and a truncate uncus. Male genitalia of both species
have been figured by Heinrich (1923).

E. separationis appears to be closely related to discretivana (Hein-
rich). Both species lack signa and sclerotization of the ductus. E. prae-
sumptiosa is thought to be most closely related to numerosana (Zeller ),
grossbecki Heinrich, abruptana (Walsingham), deflexana Heinrich, and
exacerbatricana Heinrich. This species group shares the derived char-
acteristic of long coronal setae on the cucullus. Sclerotization of the duc-
tus is also lacking in this group. MacKay (1959) included insidiosana
Heinrich in this group. However, insidiosana has short coronal setae on
the cucullus and a sclerotized band around the ductus bursae.

The distribution of separationis within the United States is limited to
southern Texas and Florida. The larvae, described by MacKay (1959),
have been reared from galls of Borrichia frutescens (L.) (Compositae )
in both localities. E. praesumptiosa is limited to southern Texas; the host
plant has not been identified.

Four Notocelia species, trimaculana (Haworth), illotana (Walsing-
ham), culminana (Walsingham), and purpurissatana (Heinrich), were
included in Epiblema by Heinrich (1923). N. suffusana, a junior subjec-
tive synonym of trimaculana, was reassigned to Notocelia from Epiblema
by Bentinck and Diakonoff (1968). MacKay (1959) described the ex-
ternal feeding larvae of culminana and trimaculana and considered them
to be a separate genus but retained them in Epiblema.

Notocelia is distinguished from Epiblema by the presence of two non-
deciduous cornuti at the apex of the aedeagus and a well-developed
lamella antevaginalis of the female. The antevaginal plate is reflexed
outward producing a projecting ostium. The ductus bursae of the female
is lightly sclerotized from the ostium to the heavily sclerotized band at
the inception of the ductus seminalis.

ACKNOWLEDGMENTS

I wish to acknowledge with my appreciation the following individuals
and institutions who have provided specimens used in this study: D. R.
Davis, U.S. National Museum of Natural History, Smithsonian Institu-
tion; J. G. Franclemont, Cornell University; Museum of Comparative
Zoology, Harvard University; W. W. Moss, Academy of Natural Sciences
of Philadelphia; E. Munroe, Canadian National Collection; F. H. Rindge,
American Museum of Natural History. Drs. Franclemont, William Mil-
ler, and Jerry Powell have read parts of the manuscript and made sug-
gestions for its improvement. I thank Amy Louise Trabka for the draw-

28 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

ings. This study was supported in part by National Science Foundation
Grant DEB 77-15808 and a Grant-in-Aid of Research from the Society of
Sigma Xi.

LITERATURE CITED

Bentinck, G. A. G. anp A. Diaxonorr. 1968. De Nederlandse Bladrollers (Tor-
tricidae). Mono. Ned. Entomol. Vereen., No. 3. 201 p., 33 pl.

Crarke, J. F. G. 1958. Catalogue of the type specimens of Microlepidoptera in
the British Museum (Natural History) described by Edward Meyrick. Vol. 3.
British Museum (Natural History), London. 600 p.

Draxonorr, A. 1950. The type specimens of certain oriental Eucosmidae and
Carposinidae. Bull. Brit. Mus. (Nat. Hist.) Ent. Vol. 1(4): 275-300, 8 pl.

_ 1973. The South Asiatic Olethreutini (Lepidoptera, Tortricidae). Zool.
Mono. Rijksmus. Nat. Hist., No. 1. i—xii, 700 p.

Forp, L. T. 1949. <A guide to the smaller British Lepidoptera. So. London Ent. and
Nat. Hist. Soc., London. 230 p.

MacKay, M. R. 1959. Larvae of the North American Olethreutidae (Lepidoptera ).
Can. Ent. 91, Suppl. 10. 338 p.

Meyrick, E. 1895. A handbook of British Lepidoptera. MacMillan and Co., Lon-
don. i-vi, 843 p.

Journal of the Lepidopterists’ Society
33(1), 1979, 28

EDITORIAL POLICY STATEMENT TO CONTRIBUTORS

Contributors are urged to pay particularly close attention to the directions for pre-
paring manuscripts for publication in the Journal. These directions are clearly spelled
out on the inside back cover of each issue. Before preparing the final copies to be
submitted, each author should decide whether or not this contribution should be in
the form of an article or a general note. Topics of articles should be of major im-
portance to the field of Lepidopterology. Please note that the format for these two
types of contributions differs. Beginning with 1 Jan. 1979, all articles should be sub-
mitted in triplicate (an original, and two reviewer's copies). Abstracts also are re-
quired for articles, but not for general notes.

Authors are urged not to submit color illustrations for publication, unless 1) they
are of exceedingly high quality, and 2) the author is prepared to pay for the cost
of these illustrations himself (the cost is approximately $550 per plate). High quality
color slides are preferred to color prints. Because of the tremendous workload in-
volving the editorial committee presently, the editor reserves the right to return those
manuscripts not conforming to the specific requirements of the Journal directly to their
authors, without review.

Austin P. PLarr, Department of Biological Sciences, University of Maryland Balti-
more County, 5401 Wilkens Avenue, Catonsville, Maryland 21228.

Journal of the Lepidopterists’ Society
33(1), 1979, 29-36

POPULATION STRUCTURE AND GENE FREQUENCY ANALYSIS
OF SIBLING SPECIES OF LETHE

MarK W. ANGEVINE AND PETER F. BRUSSARD

Section of Ecology and Systematics, Langmuir Laboratory, Cornell University,
Ithaca, New York 14853

ABSTRACT. Two recently described sibling species of Lethe (L. appalachia and
L. eurydice) exist sympatrically at McLean Bogs Reserve, Tompkins County, New
York. We examined the population structure and electrophoretic variation of these
two species and found they are substantially different. L. appalachia exhibits high
vagility in its preferred woodland habitat; delineation of spatial population units was
not feasible. L. eurydice is philopatric and local in wet meadows; demographic data
are easily obtained for this sedentary species. Interspecific comparisons of eight
enzyme-synthesizing loci revealed significant differences in allele frequencies at five
loci, providing further evidence of two separate gene pools. On the basis of these eight
loci, the calculated genetic distance between the two species is 0.145, well within the
range of values previously reported for other sibling pairs.

The existence of a pair of sibling species in the genus Lethe in eastern
North America was proposed several years ago (Cardé, Shapiro and
Clench, 1970; Shapiro and Cardé, 1970). It had long been recognized
(Field, 1936; Chermock, 1947) that at least two morphologically distinct
forms of Lethe eurydice occurred in the eastern United States. It was also
noted that most reports of this species from northern localities were from
open, wet sedge meadows. In the southern portion of the range, how-
ever, all reported individuals were observed in deep woods or bush
swamp. Chermock had placed all specimens south of Pennsylvania in
the subspecies L. e. appalachia. Cardé et al. recognized that the two
forms were, in fact, widely sympatric, habitat-isolated sibling species.
They demonstrated that a variety of subtle but consistent morphological
characters of adult genitalia and wing patterns as well as larval markings
could be used to distinguish the specimens flying in closed woods from
those appearing in open meadows. Lethe eurydice is the species char-
acteristic of open meadows. Its range extends from southern Alberta and
eastern North Dakota to Nova Scotia, and southward to Delware, Penn-
sylvania and Ohio. Lethe appalachia is restricted to shady, closed canopy
woods and bush swamp within sight of water. It is found in the Appala-
chian ranges from Georgia to Maine, and in the northern part of the range
extends westward to Michigan.

A study was undertaken at McLean Bogs Reserve, Tompkins Co., New
York to test the accuracy of Cardé and Shapiro’s analysis of the Lethe
situation. A site was chosen for sampling where putative L. eurydice
habitat was available directly adjacent to L. appalachia habitat. A mark-
recapture program was begun in the L. eurydice habitat (a very well-

30 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

marked depression with saturated soil and numerous Carex species, sur-
rounded by drier, well-grazed pasture) in order to estimate the density
of that population and measure movement of individuals into the wooded
area adjacent. A similar program was begun in the woods nearby with
L. appalachia.

Samples of individuals of each species were captured live and stored for
an electrophoretic analysis of genetic variation in the two groups, and for
a measurement of their genetic similarity.

METHODS
Mark-recapture analysis of population size

A mark, release and recapture study was undertaken in what will hence-
forth be referred to as the Pothole area to estimate the density and
turnover rate of the L. eurydice population there. The population was
sampled 14 times during the period from 5 July to 22 July 1975. Insects
were netted and secured in glassine envelopes during the 30 minute
sampling period. Afterward, each insect captured for the first time was
marked with a felt-tipped marker in a coded numerical fashion. The
specimen number, sex, and condition of all newly-captured and recap-
tured insects were recorded; and the individuals were immediately re-
leased. The point of release was changed from one day to the next to
avoid any overall displacement of individuals by capture. Most samples
were taken in the early afternoon.

The data were analyzed using the method outlined by Jolly (1965).
This method of analysis is particularly appropriate for use in studies in-
volving three or more successive samples in a population where both
dilution and loss are occurring (Southwood, 1966). The basic equa-
tion is:

P, — (Min,)/ (1)

where P, = the estimate of population size on day i, M, = the estimate of
the total number of marked animals in the population on day i, nj = the
number of individuals caught on day i, and r, = the number of previously-
marked animals caught on day i. |

Other parameters of interest estimated by the Jolly method include ¢;
the probability of survival from release time on day i to capture on day
i+], and B;, the number of new animals joining the population in the
interval from i to i+] who are still alive at time i+ 1.

Electrophoretic analysis of protein polymorphisms
Individuals of Lethe eurydice and L. appalachia were captured in the
field during the period 5 July to 22 July 1975. All 173 individuals of L.

VOLUME 33, NUMBER | 31

eurydice were collected in the Pothole area. Seventy-three individuals of
L. appalachia were collected in several areas of concentration in the
wooded areas of McLean Bogs Reserve. The boundaries of these con-
centrated areas were very diffuse, however, and individual L. appalachia
specimens were often netted in other places.

The insects were frozen live after capture and stored at -80°C until
electrophoresis. Specimens were prepared for electrophoresis by grind-
ing, after removal of legs and wings, in 0.3 ml of a pH 7.0 buffer of 0.1 mM
tris, 0.001 m EDTA and 5 x 10° m NADP. The homogenates were drawn
into capillary tubes, centrifuged at 10,000 rpm for 2 minutes and stored at
—80°C.

Horizontal starch gel electrophoresis was performed on the soluble
protein extracts by methods similar to those of Selander e¢ al. (1971). The
buffer systems used and the enzyme assays employed were as follows:
lithium hydroxide (Selander et al., 1971, buffer 2), glutamate-oxalo-
acetate transaminase (GOT), phosphohexose isomerase (PHI) and
phosphoglucomutase (PGM); continuous tris-citrate (Selander et al.,
1971, buffer 4), malate dehydrogenase (MDH, 2 loci), a-glycerophos-
phate dehydrogenase (a-GPD), and isocitrate dehydrogenase (IDH, 2
loci). A total of eight enzyme-synthesizing loci were resolved. Five of
these were polymorphic in L. eurydice, four were polymorphic in L. ap-
palachia (a polymorphic locus was defined as one in which the most com-
mon allele occurred at a frequency of >0.99). Allele frequencies were
estimated directly from the phenotype frequencies for each locus. The
formula for heterozygosity of an individual locus is H = 1-p,? where
p equals the frequency of an allele at the locus.

An estimate of the genetic distance separating the two populations was
made using the method of Nei (1975) which provides an estimate of the
mean number of codon differences per structural gene locus. This index
may take values from 0, representing populations with no alleles in com-
mon, to 1, representing populations with identical frequencies of the same
alleles.

RESULTS

Mark-recapture analysis of population size

The calculated estimates of the parameters P,, ¢, and B; for each day are

given in Table 1, and the daily estimates of population size ( P,) are
shown in Figure 1. These data indicate two distinct peaks in population
size: one on 10 July and a second, smaller peak on 14 July. Using a
method of graphical estimation (Southwood, 1966) the area under the
population estimate curve was calculated and this total, divided by the

32 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

TasBLE 1. Results of analysis of mark—recapture data.

No.

Proportion marked No. of Standard
of animals Survival new Total error of

recaptures at risk rate animals population estimation

Day A A A A A ho ae

(i) (a; ) (M;) (9;) (B;) (P,) (q Vee) )

1 — 0 158 — — 0.0
2 .136 16.6 .736 98.3 W221) TOS
3 129 24.0 IL JU) 193.3 186.0 117.0
4 nlip2 Bla) 941 330.7 378.3 Iso)
5 sll 75.8 1.68 166.1 682.9 396.4
6 .210 178.0 .296 —8.5 847.6 BE 4
7 .250 60.3 913 —15.9 2AN.2 139.0
8 382 77.0 .410 223.6 201.6 70.5
9 AS) 39.4 Wo 1F —30.9 305.4 202.3
10 Pale eee 136 28.8 MT as) eal
ith .194 (a3) — — (65.7) 1.0
12 093 (03) sa nae (32.3) 2.0
13 .066 (01) — — (15),23) 2.0
14 .000 (00) —— — — —

average adult lifetime (derived from the mean survival rate ¢;), provided
an estimate of total population size of 2,912 insects. No L. eurydice in-
dividuals were ever netted in the woods, nor were any L. appalachia
caught in the Pothole area.

A similar mark-recapture program was begun with L. appalachia at a
site in the woods about 100 meters from the L. eurydice population. This
particular site was chosen because of the consistently higher densities of
butterflies observed there in comparison to the woods in general. How-
ever, this study was abandoned because of an extremely low rate of re-
captures and a general scarcity of L. appalachia individuals. In the first
four days of the study, 29 specimens were netted, marked and released in
2 manhours of search; only one of these marked individuals was ever re-
captured. It became apparent that the site represented only a temporary
aggregation point for L. appalachia, and not a stable population unit. In-
dividual insects tended to move extensively through large areas of the
wooded bog basin, and several similar high-density sites were subsequent-
ly found.

Electrophoretic analysis of protein polymorphisms
The frequencies of all alleles for the eight enzyme-synthesizing gene
loci for each species are shown in Table 2.
In Lethe eurydice, one locus (IDH-II) was represented by a single al-
lele in all individuals. Two loci were dominated by single alleles with
frequencies greater than 99 percent (GOT and a-GPD). Two loci were

VOLUME 33, NUMBER 1 33

900

800

600

A

Pop. estimate (P; )

re

Seow 7G 9) OMe i213, 14. 15 16 17 18 19 20 21 22

Date of sample (July 1975)

Fic. 1. Daily estimates of population size (P, ) for Lethe eurydice at McLean Bogs
Reserve, Pothole area.

dominated by single alleles with frequencies greater than 95 percent but
less than 99 percent (MDH-I and MDH-II). Of the other three poly-
morphic loci, one locus had two alleles represented in the population
(IDH-I), one locus had three alleles (PHI), and one locus had four al-
leles (PGM ).

The mean heterozygosity for the population is 0.175. The heterozygosi-
ties of individual loci are listed in Table 3. The genotype frequencies at
all loci were not signifcantly different from those predicted by the Hardy-
Weinberg expression using the x? goodness-of-fit test (Sokal and Rohlf,
1969 ).

Lethe appalachia showed a similar pattern of electromorphic variation.

34 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TasLE 2. Results of analysis of allozyme variation.

Lethe eurydice Lethe appalachia
: Results of
Allele Sample Allele Sample G-test
Locus Allele frequency size frequency size (p=
GOT ar — 320 .012 162 N.S.
a id, .988 |
b .003 —
PHI a .103 340 — 164 <.001
b .679 61
c .240 A421
d — .018
PGM a orl 308 488 162 <.001 -
b 046 .265
c me, 204
d .006 043 |
MDH-I a .034 320 .006 162 : 036
b .966 .994 .
MDH-II a 975 320 1.00 162 | .O1
b .025 — |
@-GPD b 993 320 1.00 162 INES:
c .007 —
IDH-II a .876 322, .988 162 <.001
b me .O12
IDH-II a 1.00 226 1.00 150 N.S.

Three loci (MDH-II, a-GPD and IDH-II) were represented in the pop-
ulation by single alleles. One locus (MDH-1I) was strongly dominated by
an allele with a frequency greater than 99 percent. Two loci (GOT and
IDH-I) were dominated by single alleles with frequencies between 95
and 99 percent. The remaining two loci had three (PHI) and four
(PGM) alleles present in the population. |

TaBLE 3. Estimates of heterozygosity per locus.

Locus Lethe eurydice Lethe appalachia
GOT .006 024

PHI .470 DUO men
PGM 083 .648
MDH-I .066 .012
MDH-II .048 .000
a-GPD 014 .000
IDH-I pad 2 024
IDH-II .OO0 .000

Mean nus 2

VOLUME 33, NUMBER 1 BS

The mean heterozygosity of the L. appalachia pupulation was 0.152.
Heterozygosities of individual loci can be found in Table 3. No locus
showed significant deviation from Hardy-Weinberg expectation using the
x” goodness-of-fit test.

The two species are highly significantly different (p < .01, G-test) in
allele frequencies at four of the eight loci examined, and significantly
different (p = .036) at another. These data clearly indicate reproductive
isolation in sympatry. Using the genetic distance measure of Nei (1975)
the distance separating these species equals 0.145, a figure well within
the rather wide range of available estimates of distances separating sibling
pairs. (Nei’s measure can range between a maximum distance of 1.00,
representing no alleles in common, to a minimum of 0.0, representing
total identity. )

A very small number of individuals (six) of Lethe portlandia captured
at the McLean Bogs Reserve were analysed at the same eight en-
zyme loci. Although the sample size was insignificant for statistical pur-
poses, the electromorphs at six of the loci tested represented clearly dif-
ferent mobility classes from those present in either L. appalachia or L.
eurydice.

DIscuUSSION

The results of this study fully agree with the conclusion of Cardé, Sha-
piro and Clench (1970) and Shapiro and Cardé (1970) that the butter-
flies Lethe eurydice and L. appalachia are distinct, although very simi-
lar, sibling species. Both the mark-recapture study and the analysis of
genetic variation demonstrate that the two are genetically isolated and
habitat-segregated, at least in the locality studied.

The mark-recapture study indicates that the two species strongly resist
crossing over from the open to the wooded habitat, or vice versa. It also
suggests that the two populations have contrasting spatial structure.
Lethe eurydice occupies a small, isolated, concentrated, uniform patch of
acceptable habitat, while L. appalachia occupies a more “fine-grained”
habitat, moving extensively between more fragmented and diffuse sites
of maximum acceptability.

LITERATURE CITED

Carpe, R. T., A. M. SHarmo & H. K. Ctencw. 1970. Sibling species in the euryd-
ice group of Lethe (Lepidoptera: Satyridae). Psyche 77: 70-103.

CueERMock, F. G. 1947. Notes on North American Enodias (Lepidoptera). En-
tomol. News 58: 29-35.

FreLp, W. L. 1936. New North American Rhopalocera. J. Ent. Zool. (Pomona
College) 28: 17-26.

Jotty, G.M. 1965. Explicit estimates from capture-recapture data with both death
and immigration—stochastic model. Biometrika 52: 225-247.

36 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Ne1, M. 1975. Molecular population genetics and evolution. North Holland Re-
search Monographs, Frontiers of Biology, Vol. 40. North Holland Publ. Co.,
Amsterdam-American Elsevier, N.Y. and Oxford.

SELANDER, R. K., M. H. Smiru, S. Y. YANG, W. E. JoHNson & J. B. GENTRY. 1971.
Biochemical polymorphism and systematics in the genus Peromyscus. I. Varia-
tion in the old-field mouse Peromyscus polionotus. Stud. Genet. VI: 49-90
(Univ. Texas Publ. 7013).

SHapiro, A. M. & R. T. Carpe. 1970. Habitat selection and competition among
sibling species of Satyrid butterflies. Evolution 24: 48-54.

SokaL, R. R. & F. J. Rontr. 1969. Biometry. W. H. Freeman and Co., San Fran-
cisco.

SoutHwoop, T. R. E. 1966. Ecological methods, with particular reference to the
study of insects. Methuen and Co., Ltd., London.

Journal of the Lepidopterists’ Society
33(1), 1979, 36

NOTES AND NEWS

THE JAMES H. BAKER COLLECTION

The James H. Baker collection of insects has been received at the Department of
Entomology, Smithsonian Institution. Baker’s material consists of slightly more than
24,400 specimens, primarily Lepidoptera, but also contains many Coleoptera and
Diptera.

Among the Lepidoptera the collection is especially rich in Geometridae, Baker’s
specialty. Most of the specimens in this collection are from eastern Oregon, but Baker
enjoyed a wide correspondence and traded considerably; consequently there is a rather
liberal sprinkling of moths and butterflies from localities other than Baker’s home
state. Baker also collected in such places as Arizona, Idaho, and Nevada, so there is a
nice representation of species from those areas.

J. F. Gates Ciarke, Dept. of Entomology, U.S.N.M.N.H., Smithsonian Institution,
Washington, D.C. 20560. |

Journal of the Lepidopterists’ Society
33(1), 1979, 37-41

THE LARVA OF CRYPTOCALA ACADIENSIS (BETHUNE)
(NOCTUIDAE)!

TrmotHy L. McCase
New York State Museum, Albany, New York 12234

ABSTRACT. The mature larva of Cryptocala acadiensis (Beth.) (Lepidoptera:
Noctuidae) is described. Apocynum androsaemifolium LL. was found to be an ac-
ceptable food plant; additional acceptable and unacceptable food plants are listed.
Eggs were laid singly and the minimum developmental time from egg to adult was
116 days in the laboratory. This is a much shorter developmental time than what
would be expected in nature as the mature larva is presumed to overwinter.

In the type description, Cryptocala acadiensis (Bethune, 1869) was
placed in the genus Anarta Ochsenheimer because of its small size and
black bordered, yellow hind wings. Benjamin (1921) recognized it as
a synonym and the older name for Rhynchagrotis gilvipennis (Grote) and
erected Cryptocala for it. Prior to this paper nothing was known of the
life history. Mikkola and Jalas (1977) report that Rumex is the host plant
of the very close (if actually distinct) species, Noctua (Cryptocala) char-
dinyi Boisduval.

Cryptocala acadiensis occurs from Labrador south to Massachusetts and
west to the Pacific. Its flight period is from July to August (Forbes,
1954). A female of C. acadiensis was taken at ultraviolet light on 16 July
1977 in the Adirondacks, 6 mi east of Indian Lake, 1820 ft, Hamilton Co.,
New York. The following day 39 eggs were laid singly in a holding jar.
The larvae eclosed in seven days and were offered a selection of plants.

The first instar larvae initially accepted the blossoms of Hypericum
perforatum L.., Sagittaria latifolia Willd., the blossoms and leaves of Apo-
cynum androsaemifolium L., and the leaves of Prunus virginiana L., Achil-
lea millefolium L., Sambucus canadensis L., and Spiraea latifolia ( Ait.)
Borkh., but the limited feeding and continual wandering of the first instar
larvae indicated that most of these plants were unacceptable. Only A.
androsaemifolium was continuously utilized by the first instar larvae and
all later instars were reared to maturity on A. androsaemifolium leaves.
Plants refused by the first instar larvae include: Rubus idaeus L., Ame-
lanchier laevis Wieg., Pteridium aquilinum (L.) Kuhn, and Vaccinium
myrtilloides Michx.

The larvae feeding on A. androsaemifolium remained healthy (no dis-
ease) but grew slowly. They started pupating on 12 September 1977 and
adults began to emerge on 10 November 1977. Presumably, this species
would normally overwinter as a mature larva and pupate in the spring.

1 Published by permission of the Director, New York State Museum, Journal Series No. 254.

38 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fic. 1. Cryptocala acadiensis, Adirondack Mts., New York: photograph of living,
ultimate instar larva.

The larvae were cultured in tins in total darkness (interrupted only by
the addition of fresh leaves every two days) at 22°C (+8°). These
artificial conditions speeded development as is the case with many spe-
cies which normally feed at night. The early pupation is also typical of
many Lepidoptera which have a non-obligatory diapause.

The illustrations that accompany the descriptions of the last larval in-
star were drawn to scale using an ocular grid. All scale lines represent 0.5
mm. The terminology and abbreviations follow Godfrey (1972) with the
exception of coronal punctures.

General (Fig. 1). Head 2.06-2.23 mm wide. Total length 26-30 mm. Abdominal
prolegs present on third through sixth segments. Head and body smooth. Setae
simple, insertions in small, flat, black tubercles. Spiracle A-8 0.28 mm high. Seta D1
0.40 mm long.

Coloration (living material). General head and body color light brown; a light

middorsal line and a light line just below D2 on all segments; venter below spiracles —
light; spiracles also light.
_ Head (Fig. 2). Epicranial suture 1.04 mm long. Height of frons 1.40 mm. Ad-
frontal punctures (AFa) anterior and second adfrontal seta (Af-2) posterior to apex
of frons. Coronal punctures 5 (Ca-5), posterior setae 1 & 2 (Pl & P2), and lateral
seta (L.) each arise from a black pigmented spot. Ocellar interspaces between Ocl-
Oc2 and Oc2—Oc3 each equal to diameter of Oc2; Oc3—Oc4 one-third diameter of
Oc4; Oc4—Oc6 approximately 1.5 times diameter of Oc4; Oc4—Oc5 2.0 times the
diameter of Ocd4.

Mouthparts. Hypopharyngeal complex (Fig. 3): spinneret subsequal to labial
palpus, apex bearing short spinules; stipular seta (S) at anterior dorsal apex of pre-

VoLUME 33, NUMBER 1 39

Fics. 2-3. Cryptocala acadiensis, Adirondack Mts., New York: 2, frontal aspect
of head; 3, oral aspect of left mandible.

AO JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Ji Y/’
4) \ ee iMG Wij
i Y

<7) )

| Wa
Le ye
Lib if Wh Wy We

Iics. 4-6. Cryptocala acadiensis, Adirondack Mts., New York: 4, left aspect of
hypopharyngeal complex; 5, left dorsolateral setal arrangement of prothorax; 6, left
dorsolateral setal arrangement of first abdominal segment.

VoLUME 33, NUMBER 1 Al

mentum; distal region of hypopharynx covered with fine spines; proximolateral region
bearing single row of about 8 distinct spines. Mandible (Fig. 3): inner ridges dis-
tinct, with prominent basal tooth; sixth outer tooth low, divided into smaller subteeth.

Thoracic segments. Prothorax (Fig. 5): cervical shield weakly sclerotized, with
two punctures between setae D-1 and XD-1, one puncture between XD-1 and XD-2
(located two-thirds distance from XD-1 to XD-2) and one puncture along posterior
margin of shield behind D-2; SD-1 and SD-2 on same pinaculum; L1 and L2 also on
same pinaculum; SV-1 and SV-2 separate. Meso- and metathoracic segments with
a non-setiferous puncture on same pinaculum as SD-1.

Abdominal segments. Ab-] (Fig. 6): two subventral setae (SV-1 & SV-3); Ll
dorsal to spiracle. Ab-2—6 with three subventral setae. Ab-7 and Ab-8 with only one
setae in subventral group. Crochets: uniordinal, 16-17 per third abdominal proleg,
18—21 per fourth, 20-24 per fifth, 22-26 per sixth.

Material examined. Thirteen specimens, 6 miles east of Indian Lake, 1820 ft., lat.
43°45’30” long. 74°10’14”, Hamilton Co., New York, 10 September 1977, from ova of
female collected, determined, and reared by T. L. McCabe.

The basal mandibular tooth and the spinule-tipped spinneret seem to indicate a
relationship to Ochropleura plecta (L.) (see Crumb, 1956, for larval description), but
this is not substantiated by the male genitalia (figured by McDunnough, 1928).

ACKNOWLEDGMENTS

I thank Drs. J. G. Franclemont and G. L. Godfrey for reviewing this
manuscript. Mr. Stanley J. Smith verified the plant determinations and
voucher plant specimens are deposited in the New York State Museum.
Funds have been provided by the New York State Museum, Albany, New
York.

LITERATURE CITED

BENJAMIN, F. H. 1921. A study of the noctuid moths of the genera Lampra Hbn.
and Cryptocala, gen. nov. Bull. South. Calif. Acad. Sci. 20: 133.

BETHUNE, C. J. S. 1869. A new species of Anarta from Nova Scotia. Trans. Nova
Scotia Inst. Nat. Sci. 2(3): 84.

Crus, S. FE. 1956. The larvae of the Phalaenidae. U.S. Dept. Agric. Tech. Bull.
1135: 99.

Forses, W. T. M. 1954. Lepidoptera of New York and neighboring states, Cornell
Univ. Agri. Exp. Sta. Mem. 329: 71.

Goprrey, G. L. 1972. A review and reclassification of larvae of the subfamily
Hadeninae (Lepidoptera, Noctuidae) of American North of Mexico. U.S. Dept.
Agric. Tech. Bull. 1450, 265 pp.

MrKxkoLa, K. & I. Jauas. 1977. Yokkoset 1 [Noctuidae of Finland]. Otava, Hel-
singssa. 256 pp.

McDunnoucu, J. H. 1928. A generic revision of North American Agrotid moths.
Nat. Mus. Canada Bull. 55, 78 pp.

Journal of the Lepidopterists’ Society
33(1), 1979, 42-49

COURTSHIP BEHAVIOR OF THE CHECKERED WHITE, PIERIS
PROTODICE (PIERIDAE)

RonALD L. Rutowski
Department of Zoology, Arizona State University, Tempe, Arizona 85281

ABSTRACT. Courtship behavior leading to copulation is described for the check-
ered white, Pieris protodice, from film records of courtships with perched virgin fe-
males and written records of courtships elicited by releasing virgin females near free-
flying males. Temporal and sequential patterns of successful courtships follow those
seen in other pierid butterflies in that the interactions are simple and rapid, averaging
about 3 seconds in duration. Unsuccessful courtships are also described and conform
to patterns documented for other pierids. The checkered white differs from con-
familials only in that there is no abdominal extension by females during successful
courtship and after copulation a post-nuptial flight may occur. The possible func-
tions of post-nuptial flights are discussed.

Although ethologists have been interested in butterfly courtship for
many years (Scott, 1974; Silberglied, 1975) few quantitative studies of
the temporal and sequential structure of butterfly courtship can be found
in the literature. Detailed information is essential for comparative studies
of butterfly courtship which in the past have proven useful in deducing
the ecological factors influencing the structure of both successful and
unsuccessful courtships (Brower et al., 1965; Pliske, 1975; Rutowski,
1978a).

In this paper the courtship of the checkered white, Pieris protodice
Boisduval and Le Conte, is described in detail. Attention will be focused
on the structure of successful courtships so that this account can be com-
pared to previous studies of a similar nature, with special reference to
the function of post-nuptial flights in butterflies. It should be noted that
a brief description of P. protodice courtship was given in Abbott (1959).
However, Shapiro (1970) has pointed out the interpretational problems
surrounding that description, so it will not be dealt with here.

METHODS

All observations were made and all animals obtained from March
through June in 1976 and 1977 at the Arizona State University Field Lab-
oratory, Tempe, Arizona. To obtain virgin females, eggs were collected
by placing field-caught females in tubular cheesecloth cages (1 m high,
0.25 m diameter) with cuttings of the local larval foodplant, Sisymbrium
irio L.. The larvae from these eggs were reared to adulthood in the labora-

tory on S. irio. The humidity and light regimen in the rearing area were
not regulated and variable.

VoLUME 33, NUMBER 1 43

All behavioral observations were made on clear days between 0900
and 1500 when the butterflies were most active. Naturally-occurring in-
teractions between males and between males and females were observed
and the form, outcome, and duration (as timed with a stopwatch) of each
recorded. Similar records were made of interactions initiated by releasing
virgins near free-flying males.

During late May 1977, films of successful courtships were made at 24
and 70 frames per second using a Beaulieu 4008 ZM II super-8 movie
camera. In all cases a virgin female was placed near the top of an ex-
posed perch in a large patch of S. irio where there was a dense popula-
tion of flying males. By activating the camera as a male approached a
complete record of the courtship could be obtained. Temporal and se-
quential data were gathered via frame-by-frame analysis of the film
records.

Where pertinent, summary statistics are given as mean + standard
error of the mean.

RESULTS
I. Successful Courtship

Using 38 virgin females, 27 successful courtships (= ending in copula-
tion) were recorded on film and 58 were observed after releasing virgin
females near free-flying males. Temporal data from the films will be
summarized first to give a general impression of the structure of P. proto-
dice courtship.

A. Film records

Successful courtship with a perched female began when the male's
wings or legs made physical contact with the female. In 67 percent of the
filmed courtships this contact was made with the legs as the male alit on
the female and immediately walked toward her thorax. In all other film
records the male broke and then renewed contact with the female at least
once, and as many as four times, before positioning himself on the female’s
thorax. Once on the female's thorax in a head-to-head orientation the
male curled his abdomen out from between his hindwings and inserted
the tip between the female’s hindwings, often after several unsuccessful
attempts. It was not possible to determine when genital contact was
made. The courtship ended when the male stopped moving his wings
and assumed a quiescent posture. Males showed no preference for the
side of the female from which they effected copulation (13 right vs. 14
lest — 0.057, p — 0:8).

Fig. 1 shows the temporal pattern of the following major events in a
successful courtship with a perched female: male contacts female, male

44 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

contacts )

oO begins probing(n=13)

S| abdomen between ) wings (n=6)

& stops moving (n= 27)

a
—— i SS aie
O 0.5 1.0 1.5 20 22 3.0 3.5 40 125

SECONDS AFTER BEGINNING OF COURTSHIP (=c'CONTACTS 9)

Fic. 1. Temporal relationships between the major behavioral events in the success-
ful courtship of P. protodice. The mean time of occurrence, standard error (open bar),
range line, and sample size (n) are shown for each behavior. See text for details.

begins probing (abdomen appears from between hindwings), male ab-
domen between female hindwings, male stops moving. There were no
observed variations in the sequence of these events. In 45 percent of the
courtships the female performed a low amplitude flutter response. Of
these, 25 percent began before the male first contact and 75 percent after,
and they ended most often before the male began probing but sometimes
not until he had inserted his abdomen between the female’s hindwings.
Detailed quantitative information on these points was difficult to gather
because of 1) the viewing angle of many of the film records and 2) the
low amplitude of the flutter response. In any event, flutter responses had
no significant effect on the duration of courtship (courtships with flutter
response: 2.8 + 0.37 sec, n = 12; courtships without flutter response: 3.86
+ 0.93 sec, n = 15; t = 0.968, p = 0.66).

Shortly after coupling some males attempted to fly away from the perch
carrying the female. In the 11 courtships where these attempts were
filmed the males broke contact with the perch on the average 3.71 + 0.586
sec (range = 1.71-7.34 sec) after they had stopped moving their wings.
In 8 of these attempted “post-nuptial flights” (Brower et al., 1965) the
female did not release her grip on the perch and the male dangled from
the female’s abdomen at the end of the attempt. In the other 3 cases the
male flew off carrying the female to a distant and usually less exposed
perch. Systematic data on these points or on the duration of successful
post-nuptial flights was not collected.

VoLuME 33, NUMBER 1 45

wl? i

Ss Q Tying

a 8 Mmedian=7 sec

es a= SxS)

=/6

O

U4

5

ieee

2 Bib peere|n

Y)

lid] 10)

O

O

Ea)

VY)

Le perched

O median=6 sec

aia n=9

A

3 ele

2 6 an on
Mic. Sac... 2
eee

DURATION (SEC)

Fic. 2. A frequency histogram of the duration of successful courtships with
perched and flying females. Courtships were elicited by releasing females near free-
flying males and were timed with a stopwatch. n = number of courtships.

B. Written records

The successful courtships elicited by releasing virgin females near free-
flying males were timed beginning when the two animals arrived within
1 to 2 cm of each other. The distance at which these observations were
made did not allow the use of contact as a criterion for when courtship
began. This coupled with the lack of precision in determining when
courtship ended resulted in the apparent greater length of these court-
ships relative to those timed from film records. These courtships were
divided into two groups depending on whether the female was flying or
perched when the male arrived within 1 to 2 cm of her (Fig. 2). The dif-
ferences between these two types of courtships in the distribution of
durations and median durations were slight suggesting that aerial com-
ponents were of short duration. In fact, flying females typically landed
immediately when approached by a male. The form and sequence of all

46 JoURNAL OF THE LEPIDOPTERISTS SOCIETY

ground components of these courtships were exactly like those described
from the film records.

Post-nuptial flights were attempted in 27 of the 58 successful courtships
with released females. This should not be taken as an absolute indicator
of the frequency of attempted post-nuptial flights because no firm proto-
col was established for how long a copulating pair should be watched
after the courtship to wait for the onset of a post-nuptial flight attempt.
Of the observed attempts 9 were unsuccessful, i.e., the female did not re-
lease her grip on the perch where copulation occurred.

The duration of copulation was timed for 15 pairs. Thirteen of these
pairs copulated for less than 35 min (27.2 +1.17 min). The other two
pairs copulated for 179 and 236 minutes, respectively.

II. Unsuccessful Courtships

Interactions between males and virgin females that did not result in
copulation are here referred to as unsuccessful courtships. Film records of
unsuccessful courtships were not made because of the great variation in
their form and duration. Data on the following five general types were
derived from written records of 34 unsuccessful courtships between males
and released virgins. The number of observations (n) and the percent
of all documented unsuccessful courtships is given for each type. All types
terminated when the male departed.

Type A. The female continued flying on a level course (n = 17, 50%). These
interactions were of short duration (2 to 15 sec). The males were probably of low
courtship persistence like those seen by Rutowski (1978b) in Colias butterflies.

Type B. The female continued flying and initiated an ascending flight (n = 1,
3%). As well as the one interaction with a virgin female, seven naturally-occurring
interactions were seen in which the females (presumably mated, see Shapiro, 1970)
initiated ascending flights when courted by males. The interactions ranged in dura-
tion from 5 to more than 35 sec. Since all observed ascending flights were between
males and females and since none ended in copulation it is assumed that they were
attempts by mated females to curtail the courtship of persistent males as has been
documented for the ascending flights of Colias butterflies (Rutowski, 1978b).

Type C. The female alit on vegetation or on the ground; the male may or may not
have attempted copulation (n= 7, 20%). Since these unsuccessful courtships ranged
in duration from 3 to 10 sec, the males were probably of low persistence as in type A.

Type D. The female alit on vegetation or on the ground and performed a flutter
response, a pierid mate refusal posture, or both (n =9, 26%). These interactions
were from 10 to 68 sec in duration. When perched females spread their wings and
elevated the abdomen they were said to have performed the mate refusal posture first
described by Obara (1964) for P. rapae and reported for several other pierids (Scott,
1973). Abbott (1959) previously and incorrectly descibed the mate-refusal posture
as an immediate invitation to copulation in P. protodice. This posture mechanically
impedes the male’s copulatory attempts. Females achieved the same effect in other
situations by performing the flutter response, a rapid opening and closing of the wings
(Obara, 1964; Rutowski, 1978a).

Type E. The male was displaced by another male (n= 1, 3%). This was only
seen once with a virgin although I have also seen it occur with wild females. A

VoLUME 33, NUMBER 1 47

courting male terminated his courtship attempt when another male approached the
female and began courtship. The second male was also unsuccessful. The female in
this case was perched during the displacement. In naturally occurring unsuccessful
courtships two males courting the same perched or flying female may leave together
while circling each other in rapid flight.

Systematic observations of unsuccessful courtships with mated females
were not made but casual observation suggests that they may only be of
type A, B, D, or E. Shapiro (1970) noted that such courtships may be

very lengthy in duration, up to 30 min or more.

DISCUSSION

Successful courtship in Pieris protodice is rapid and highly stereotyped.
There are no prolonged aerial components and once the female alights on
vegetation or on the ground the male does little more than land on her
thorax and couple with her. This description closely fits that given for
the temporal and sequential characteristics of the courtships of other
pierids including Eurema lisa Boisduval (Rutowski, 1978a), Colias eury-
theme Boisduval, C. philodice Latreille (Silberglied and Taylor, 1978),
and four species of Pieris in Japan (Suzuki et al., 1977). Pieris protodice
also shares components of its courtship with Leptidea synapsis Linnaeus
although the latter's courtship is apparently longer and includes some
striking male displays (Wiklund, 1977). However, the most obvious dif-
ference between the courtship of species in the genus Pieris and that of
other pierids is the lack of an abdominal extension response on the part
of the female. In E. lisa, L. synapsis, and Colias species the male cannot
couple with the female unless she extends her abdomen ventrally out from
between the hindwings. Chemical and tactile cues delivered as the male
courts the female elicit this response in E. lisa females (Rutowski, 1977).
A male of P. protodice must insert his abdomen between the female’s hind-
wings to reach her abdomen and couple. At present the proximate and
ultimate causes of this variation are unknown.

The unsuccessful courtships of P. protodice also follow patterns ob-
served in other pierids. Males vary in persistence and females utilize mate
refusal postures, flutter responses, and ascending flights to curtail or im-
pede the copulatory attempts of persistent males. Interestingly, all three
behavior patterns were displayed by virgins as well as mated females sug-
gesting that males vary in their attractiveness to virgin females which may
be selective in their choice of a mating partner. Similar responses to males
by virgin E. lisa females have been hypothesized to serve the same func-
tion (Rutowski, 1978a). The rejection responses of virgin Colias females
definitely play a role in avoiding courtships with males of the wrong
species (Taylor, 1973).

Post-nuptial flights have been previously reported only for danaids by

48 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Brower et al. (1965) and Pliske (1975). It should be noted that these
post-nuptial flights are spontaneous and as such are distinguished from
the induced flights of copulating pairs summarized in Shields and Em-
mel (1973). To date the only function attributed to post-nuptial flights
is that of removing the copulating pair from the place where courtship ac-
tivity has made them conspicuous to one where they are less visible
(Brower et al., 1965). In P. protodice courtships, females often land on
exposed perches. Presumably this facilitates copulation since males fre-
quently appear to have trouble locating and mating with females that
land in dense vegetation. Thus, post-nuptial flights in this species may
also serve an anti-predator function. As an extension of this hypothesis,
post-nuptial flights may also remove animals from exposed perches where
thermal stress might become a problem, especially on warm days.

The selective pressures of predation and thermal stress should act
equally on copulating males and females. However, the data clearly in-
dicate that males and females are not equally willing to participate in
post-nuptial flights. I suggest that there may be conflict between males
and females as a result of male—male competition. Most courtships occur
in or near stands of larval foodplant where males look for receptive ovi-
positing females or newly-eclosed virgin females. During copulation, pairs
are often buffeted by single males in a way that makes them conspicuous
to potential predators or may result in the separation of the pair. From
the male’s perspective both would be detrimental to his fitness and select
for copulating males who initiate post-nuptial flights and move to areas of
low male density. However, from the female’s perspective it may be ad-
vantageous to remain in the vicinity of a stand of larval foodplant for the
following reason. Because males do not defend resources of interest to
females, females may select males only on the basis of behavioral cues
that are correlated with high genetic quality. One of these cues may be
the male’s ability to defend the pair against interruptions by other males
during copulation. If so, then it may be to the female’s advantage to stay
in an area of high male density thereby forcing the male to fend off the
onslaughts of other males. If he cannot, then the female might benefit by
copulating with another male.

In summary, I propose that selection does not act equally on males and
females with respect to their participation in post-nuptial flights. In par-
ticular, there appear to be ecological circumstances in which males bene-
fit from post-nuptial flights but at least some females do not because the
potential benefits of preventing post-nuptial flights and inciting male-
male competition outweigh the potential costs of increased predation and

VOLUME 33, NUMBER 1 49

thermal stress. The ecological conditions that give rise to this situation
remain to be defined.

ACKNOWLEDGMENTS

I thank Dr. John Alcock for his critical reading of the manuscript and
Mr. Larry Marshall for his assistance in the field. This study was sup-
ported in part by an Arizona State University Faculty Grant-in-Aid and
National Science Foundation Grant BNS 78-11211.

LITERATURE CITED

AspspotT, W. 1959. Local autecology and behavior in Pieris protodice Boisduval
and Leconte with some comparisons to Colias eurytheme Boisduval. Wasmann J.
Biology 17; 279-298.

Brower, L. P., J. V. Z. BRowEer & F. P. Cranston. 1965. Courtship behavior of
the queen butterfly. Zoologica (N.Y.) 50: 1-39.

Oxpars, Y. 1964. Mating behavior of the cabbage white, Pieris rapae crucivora. II.
The ‘mate-refusal posture’ of the female. Dobut. Zasshi 73: 175-178.

Opara, Y. & T. Hipaxa. 1964. Mating behavior of the cabbage white, Pieris rapae
crucivora. I. The ‘flutter response’ of the resting male to flying males. Dobut.
Aassmi(o- V31—135.

PutskE, T. 1975. Courtship behavior of the monarch butterfly, Danaus plexippus
L. Ann. Entomol. Soc. Amer. 68: 143-151.

Rurowsk1, R. L. 1977. Chemical communication in the courtship of the small sul-
phur butterfly Eurema lisa (Lepidoptera, Pieridae). J. Comp. Physiol. 115:
75-85.

1978a. The courtship behavior of the small sulphur butterfly Eurema lisa

(Lepidoptera, Pieridae). Anim. Behav. 26: 892-903.

. 1978b. The form and function of ascending flights in Colias butterflies.
Behav. Ecol. Sociobiol. 3: 163-172.

Scott, J. A. 1973. Mating of butterflies. J. Res. Lepid. 11: 99-127.

SHAPIRO, A. M. 1970. The role of sexual behavior in density-related dispersal of

pierid butterflies. Am. Nat. 104: 367-372.

SHIELDs, O. & J. F. EMMEL. 1973. A review of carrying pair behavior and mating
times in butterflies. J. Res. Lepid. 12: 25-64.

SILBERGLIED, R. E. 1977. Communication in the Lepidoptera. In: How Animals
Communicate (ed. T. Sebeok). Indiana University Press, Bloomington.

SILBERGLIED, R. E. & O. R. Taytor, Jr. 1978. Ultraviolet reflection and its be-
havioral role in the courtship of the sulphur butterflies Colias eurytheme and C.
philodice (Lepidoptera, Pieridae). Behav. Ecol. Sociobiol. 3: 203-243.

Suzuki, Y., A. NAKxaAnisni, H. Sura, O. Yata & T. Satcusa. 1977. Mating be-
haviour in four Japanese species of the genus Pieris (Lepidoptera, Pieridae).
Kontyu 45: 300-313.

Taytor, O. R., JR. 1973. Reproductive isolation in Colias eurytheme and C. philo-
dice (Lepidoptera: Pieridae): Use of olfaction in mate selection. Ann. En-
tomol. Soc. Amer. 66: 621-626.

Wrixtunp, C. 1977. Courtship behaviour in relation to female monogamy in Lep-
tidea synapsis (Lepidoptera). Oikos 29: 275-283.

Journal of the Lepidopterists’ Society

33(1), 1979, 50-55

A NEW TECHNIQUE FOR THE PROSPECTIVE SURVEY OF SEX
CHROMATIN USING THE LARVAE OF LEPIDOPTERA

WINIFRED Cross AND ALISON GILL

Department of Genetics, University of Liverpool, P.O. Box 147, Liverpool, L69 3BX,
WRK

ABSTRACT. A way of examining the heteropyknotic body is described, using cells
from prolegs amputated from living larvae. Larval survival rate is high and the results
are accurate. Prospective testing for the presence or absence of sex chromatin is par-
ticularly valuable in studying intersexes, e.g., in Lymantria dispar, where the adult
phenotype is not necessarily an indication of the chromosome constitution of the larva.

It is well known that in Lepidoptera a heteropyknotic body may be
found in the somatic cells of the female whereas it is lacking in the male
(Smith, 1945). However, there are some exceptions to this rule (Traut
and Mosbacher, 1968); for example, in Papilio machaon L. the male is
polymorphic for the character (Clarke et al., 1977). There is also good evi-
dence that in the female the body is derived from the W(=Y) chromo-
some (Suomalainen, 1969; Traut and Rathjens, 1973) and that where it
is present in the male it is associated with a particular autosome (Clarke
et al., 1977).

Testing for the “Smith” status has usually been carried out on freshly
killed larvae or adults, but Daker (1977) showed that in Hypolimnas bo-
lina L. it was possible to assess it from a spine taken from a living larva
which thereafter usually developed normally.

In the present paper we show that it is also possible to obtain good
preparations using a proleg of last instar larvae. We, in fact, found that
our preparations from prolegs were of better quality than those from
spines. Moreover, this method is particularly useful in dealing with lar-
vae which have no spines.

MATERIALS AND METHODS

The species investigated were Papilio glaucus L., Papilio dardanus
Brown, Ewploea core amymone Cr., Hypolimnas bolina L. and Lymantria
dispar L.

The P. glaucus stock was bred at Caldy, Wirral, and derived from two
females from Virginia, U.S.A., kindly supplied by Prof. J. J. Murray. Mrs.
Jennifer Maddison of Ibadan sent the butterflies from which all the
Nigerian P. dardanus stock was bred and Mrs. Gweneth Johnston posted
to us living E. core butterflies from Hong Kong which produced the
tested larvae. In H. bolina, race hybrid stock was used, the parent forms
coming from Sarawak (from Mr. Stephen Kueh) and Sri Lanka (from Mr.

VoLUME 33, NUMBER 1 51

Fic. 1. Proleg tip of H. bolina larva showing tissue scraped out. (As seen under
dissecting microscope. )

Fic. 2. Nuclei of tissue cells containing heteropyknotic body from a larva which
developed into a female butterfly. (Using x90—oil immersion—objective. )

52 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fic. 3. Nuclei of tissue cells lacking heteropyknotic body from a larva which de-
veloped into a male butterfly. (Using x90—oil immersion—objective.) Editor's
note: photograph reduced to *4 size of original.

P. B. Karunaratne. The pure Japanese broods of L. dispar originated from
a wild Nagoya female supplied by Dr. Shigeru Ae, and the hybrid brood
was from a mating between a German female (from Herr Willy Schultz)
and a bred Nagoya male.

In the early experiments, the larva to be tested was lightly anesthetized,
and then the extreme tip of one of the abdominal prolegs (Fig. 1) was
removed with a sharp pair of dissecting scissors. Later it was found that
better survival was obtained without an anesthetic. Enough tissue can be
scraped from the inside of the proleg to make one good preparation; the
material is teased out and spread as thinly as possible. After the amputa-
tion each larva was kept separately.

The cells are not fixed before staining. Two drops of 2% orcein in 457%
acetic acid are placed over the tissue and a coverslip added immediately.
After 10-15 minutes the coverslip is firmly pressed to make a “squash”
preparation. The “Smith” body when present can be clearly seen under

the 40 objective as well as under the <90 (oil immersion) objective
(Figs. 2&3).

RESULTS

Results are shown in Table 1. The accuracy of the method is assessed
by noting the sex of the butterfly or moth when it emerges, and in the

53

VoLuME 33, NUMBER |

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‘OATILSOU . YPIUIG,, 919M PUL DAT}

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A][eu1e}xe poxos sosvys [ednd » BAILY Po}So} OVAILT Apoq ,,q}1urg,, sotoeds » ‘ou poolg

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54 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

case of overwintering insects, by scoring the sex of the pupa by its external
appearance. Concordance between the larval score and the adult or pupal

sex is high.

DiIscussION

Several points are of interest:

In P. glaucus, it had been reported previously (Clarke et al., 1976) that
the black females were “Smith” positive and the yellow ones (which are
male-like) and the males were negative. It is now clear that frequently
yellow females are positive, and there appears to be a polymorphism for
the character in the yellow form. All black females have, however, so far
been positive.

In P. dardanus, in the present material, the hippocoon females are con-
sistently negative, though previously a few insects of this and other fe-
male forms have been positive, so that here again there is a polymorphism.

This information in both these species could clearly have been obtained
without sexing the larva, but prospective testing has the great advantage
that it is possible to select and breed from a female of known “Smith”
status. Moreover, it obviates the necessity of testing her immediately
after death which is obligatory because rapid degeneration of the cell
nuclei occurs post-mertem.

The most valuable application of the method, however, will become
evident when L. dispar is further studied. Here, in race crosses, inter-
sexes may occur (Goldschmidt, 1933), and it will be most informative to
relate the sex chromatin status to the phenotype and to the gonadal mor-
phology.

ACKNOWLEDGMENTS

We are extremely grateful to Sir Cyril Clarke F.R.S. for his help in
writing this paper and for the use of his facilities and livestock, to Mr.
Maurice Gill for taking the photographs and to all those named in the text
who kindly provided the living material.

LITERATURE CITED

Ciarke, C. A., P. M. SHepparp & U. Mirrwocu. 1976. Heterochromatin poly-
morphism and colour pattern in the tiger swallowtail butterfly Papilio glaucus L.
Nature, Lond. 263: 585-587.

CLanke, C, A., U. Mirrwocu & W. Traur. 1977. Linkage and cytogenetic studies
in the swallowtail butterflies Papilio polyxenes Fab. and Papilio machaon L. and
their hybrids. Phil. Trans. R. Soc. Lond. B 198: 385-399.

Daker, M. G. 1977. Report of R.E.S. Meeting. Antenna, Bull. R. Entomol.
Soc. Lond. 1: 22

GoLpscumipr, R. 1933. Lymantria. Bibliographia Genetica 11: 1-186.

VOLUME 33, NUMBER 1 Do

SmirH, S. G. 1945. The diagnosis of sex by means of heteropycnosis. Sci. Agric.
25: 566-571.

SUOMALAINEN, E. 1969. On the sex chromosome trivalent in some Lepidoptera
females. Chromosoma ( Berl.) 28: 298-308.

Traut, W. & G. C. Mospacuer. 1968. Geschlechtschromatin bei Lepidopteren.
Chromosoma ( Berl.) 25: 343-356.

Traut, W. & B. Ratyyens. 1973. Das W-Chromosom von Ephestia kuehniella
(Lepidoptera) und die Ableitung des Geschlechtschromatins. Chromosoma
(Berl. ) 41: 437-446.

Journal of the Lepidopterists’ Society
33(1), 1979, 55

BOOK REVIEW

BurrERFLIES. Text by Jo Brewer, photographs by Kjell B. Sandved. 1976. Harry N.
Abrams, New York. 176 pp., ill. Price: hardcover, $18.95; softcover, $9.95, U.S.

In the last few years a number of fine popular volumes with exquisitely colored
plates have been published, but none has had such an innovative and refreshing ap-
proach as this book. Since it is broad in scope and supplies the necessary basic in-
formation for the study of Lepidoptera in the clear, concise manner for which Brewer
is duly noted, the book should stimulate an interest in and appreciation for the insect
group from a technical as well as an aesthetic point of view. It is well illustrated
with 245 photographs (133 in color) and additional line drawings and scanning elec-
tron micrographs.

The organization of the book is quite a departure from traditional treatments. There
is a section on the economic impact of butterflies on man (“Historical Notes on
Butterflies, Moths and Men”). The section on “Butterflies in Art, Heraldry and
Religion” which chronicles the symbolic impact of butterflies on man in everyday life
and in legend is especially noteworthy. The remaining sections delve into those areas
which man finds so curiously fascinating: metamorphosis, ornamentation of the
wings, the compound eye and protective devices. The section on the wings not only
examines the physical aspects in terms of wing scales, pigmentation and wing forma-
tion, but also the mechanics involved in temperature regulation and flight, all through
the enchanted photographic eye of Kjell Sandved. In “Protective Devices,” decep-
tion, warning coloration and camouflage are discussed. There is also a brief explana-
tion of Batesian and Miillerian mimicry, along with a discussion of larval specificity
on certain toxic hostplants and the important role which these plants play in mimetic
associations.

In such a volume which includes an array of photographs, there are some or-
ganizational problems in fitting the plates with the appropriate text. The last 25
pages illustrate further intricate designs and structural iridescence, so intriguing to
the natural observer. While these are interesting, they seem somewhat superfluous.
In a few cases the identifications are incorrect or not in keeping with current litera-
ture such as Thecla syncellus (=Panthiades bitias ).

The above points by no means diminish the utility and significance of this book
for its intended audience. Its true value will be realized indeed by the enthusiasm
and appreciation generated for this diverse biological group in both aspiring and pro-
fessional lepidopterists alike.

JACQUELINE Y. MILLER, Allyn Museum of Entomology, 3710 Bay Shore Road,
Sarasota, Florida 33580.

Journal of the Lepidopterists’ Society
33(1), 1979, 56-57

GENERAL NOTES

OVIPOSITION OF THE BUTTERFLY BATTUS BELUS VARUS
(PAPILIONIDAE )

Members of the swallowtail genus Battus use as larval foodplants woody vines in the
family Aristolochiaceae (Brower & Brower 1964, Zoologica 49: 137-159; Ehrlich &
Raven 1965, Evolution 18: 586-608; Young 1971, Rev. Biol. Trop. 19: 210-240;
Tyler 1975, The Swallowtail Butterflies, Naturegraph, California). The Aristolochia-
ceae, like many vines, reach their greatest diversity in the New World tropics (Pfeifer
1966, Ann. Missouri Bot. Garden 53: 1-114), where both Battus and the closely
related Parides exploit them. Although the neotropical B. polydamas carefully de-
posits small clusters of eggs on various species of Aristolochia in Costa Rica (Young,
op. cit.; pers. obs. ), the temperate zone B. philenor is known to oviposit on plants other
than Aristolochiaceae, which are not acceptable to larvae (Tyler, op. cit.). Parides
always deposits eggs precisely on Aristolochiaceae (Young 1973, Psyche 80: 1-21;
1976, J. Lep. Soc. 31: 100-108). Battus belus varus Kollar ranges from Vera Cruz,
Mexico to northeastern Ecuador and northern Venezuela (Rothschild & Jordan, 1906,

Fic. 1; _ Battus belus varus ovipositing on Melothria guadalupensis (Curcubita-
ceae ) at Minca La Tigra, near La Virgen de Sarapiqui, Heredia Prov., Costa Rica, 19
February 1977, 1300 hrs.

VoLUME 33, NuMBER 1 57

Noyitates Zool. 13: 27-753; Tyler, op. cit.); larval foodplants are various species of
Aristolochia (e.g., Tyler, op. cit.). This note reports an observation of oviposition of
B. belus varus on a plant other than a larval foodplant.

On 19 February 1977, a female B. belus varus was observed flying among several
clumps of woody and herbaceous vines in a secondary forest at Finca La Tigra, near
La Virgen de Sarapiqui, Heredia Province, Costa Rica. She finally began ovipositing
on a vine, Melothria guadelupensis (Spreng.) Cogn. (Cucurbitaceae), intertwined
with another vine, Aristolochia constricta Griseb.; the leaves of the two plants were
similar in size and general shape. Oviposition lasted several minutes (Fig. 1). Forty-
three eggs were deposited in a tight cluster on a single leaf of M. guadelupensis and
no eggs were found on the A. constricta. In the laboratory, the freshly hatched larvae
did not accept leaves of M. guadelupensis, but fed briefly on A. ringens Vahl (ob-
tained from H. W. Pfeifer in 1971; locality not specified) before dying. Aristolochia
constricta was not available for testing.

Tyler (op. cit.) mentions that B. philenor accepts only certain species of Aristolo-
chiaceae as foodplants. Foodplant specificity is apparent where different species of
Aristolochia occur in the same region (Scriber & Feeny 1976, J. Lep. Soc. 30: 70-71).
An Australian Aristolochia-feeding swallowtail, Ornithoptera priamus, has been ob-
served to deposit eggs on an introduced species of Aristolochia, and the larvae perished
(Straatman 1962, J. Lep. Soc. 16: 99-103). The refusal of A. ringens, a plant spe-
cies native to Costa Rica, by B. belus larvae, supports the possibility that Battus
specializes on restricted larval foodplants within the Aristolochiaceae. Eggs of B.
vhilenor have been found on Convolvulaceae and Polygonaceae, vines which generally
look like Aristolochia. Larvae of another Aristolochia-feeding swallowtail, Polydorus
aristolochiae (Fabricius ), have been seen on various Cucurbitaceae in India, but their
larvae refused to accept these plants in captivity (Ghosh 1914, Mem. Dept. Agr.
India, Entomol. Sec. V(1): 53-587). Only certain species of Aristolochia are food-
plants of P. aristolochiae (Munshi & Moiz 1967, J. Lep. Soc. 21: 127-128). It is pos-
sible that B. belus varus mistook the cucurbit vine for an Aristolochia. Perhaps the
very close proximity of the A. constricta vine contributed to this confusion, by provid-
ing odoriferous and visual properties of a correct foodplant. Alternatively, the ovi-
position on the intertwined cucurbit might have been deliberate, possibly representing
an adaptation to avoid waiting egg parasites and predators. Under this explanation,
the newly hatched larvae would have rapidly found the correct foodplant. Further
observations are needed to distinguish between these two hypotheses. If an adaptation
for avoiding egg parasites and predators, such behavior might be more prevalent
among vine-feeding butterflies in the tropics, where the intertwining of unrelated vines
is common.

ALLEN M. Younc, Invertebrate Division, Milwaukee Public Museum, Milwaukee,
Wisconsin 53233.

Journal of the Lepidopterists’ Society
33(1), 1979, 57-58

MALFUNCTION OF ECDYSIS ALLOWING IMAGINAL EMERGENCE
BUT CAUSING DEATH OF ADULT HACKBERRY BUTTERFLY
(NYMPHALIDAE)

Insects must periodically shed their skins—a process known as ecdysis which al-
lows growth or transformation of the individual. Each molt period is a dangerous
time during which the insect is susceptible to predation or physiological malfunc-
tioning, both of which may cause death. Natural selection has, therefore, perfected
the process of ecdysis to such a degree that physiological failures are rare. I describe

58 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

below the partial failure of an act of ecdysis—a developmental malfunction which did
not prevent transformation, but one that was eventually fatal to the butterfly involved.
Such malfunctions are not uncommon in laboratory cultures; however, this observation
is of interest because it involved a wild-caught adult.

On 17 July 1977 at 1100 hours CDT, I was handed an adult female Asterocampa
celtis antonia (Edwards) which had been hand-collected in a residential backyard
in Austin, Travis County, Texas. Lack of worn spots on the fully expanded wings,
occasional release of untransformed fluids and an egg-packed abdomen indicated
recent emergence, probably that same morning. I soon discovered that the head was
covered with partial exuviae which had not been properly shed. Practically the en-
tire chrysalid head capsule was still present covering the greater part of the imaginal
head. Additionally, the left side of the larval head capsule was still attached to the
outside of the chrysalid head capsule.

Upon metamorphosis of the prepupa to pupa, the left half of the final-instar larval
head capsule failed to separate from the newly formed pupal epidermis. When ima-
ginal emergence occurred, both sides of the pupal “head” failed to separate as a result
of mechanical restriction caused by the still-present larval head capsule section.

Although the adult was able to emerge and properly expand and dry its wings,
damage to selected parts of the head effectively negated any chance that this in-
dividual would reproduce. Both eyes appeared completely normal and allowed reac-
tion to approaching objects. The left antenna was not visible, having been trans-
formed into an unrecognizable mass associated with the remnant exuviae. The right
antenna was free and fully developed; however, it was flexed laterally and horizontal-
ly (about 75° from perpendicular) about 2 mm from its base. This antenna could
be moved at its base, but such movements occurred only when the antenna was
touched; no spontaneous movements were observed. The most significant damage
affecting the fitness of this individual involved its proboscis. The proboscis was en-
tirely nonfunctional because of failure of the two maxillae to properly fuse. The two
halves adhered to each other in a haphazard manner and to the remaining exuviae; re-
moval of the exuviae was accompanied by removal of the proboscis halves.

Lack of a functional proboscis caused early death of this individual, because the
butterfly was unable to feed or obtain moisture. This damaged individual grew pro-
gressively weaker until it died approximately 75 hours following capture (in a cage at
an ambient diurnal temperature range of about 25-35°C). I thank Patrick K. Neck
for supplying the specimen.

RayMonp W. Neck, Pesquezo Museum of Natural History, 6803 Esther, Austin,
Texas 78752.

Journal of the Lepidopterists’ Society
33(1), 1979, 58-60

AGGREGATIVE BEHAVIOR OF ANARTIA FATIMA (NYMPHALIDAE) IN-
GUANACASTE PROVINCE, COSTA RICA DURING THE DRY SEASON

The neotropical butterfly Anartia fatima Fabricius (Nymphalidae: Nymphalinae )
is widespread throughout the coastal wet and dry regions of Central America and
northern South America (Godman & Salvin 1879, Biologia Centraliamericana, Insecta,
Lepidoptera-Rhopalocera, vol. 1, 487 p.). Several larval foodplants, in the Acanthaceae,
are shared with other nymphalines such as Siproeta (Young & Muyshondt 1974, Stud.
Neotrop. Fauna 9: 155-176), but during the severe dry season of dry regions these
plants exhibit leaf drop and become unsuitable for oviposition (Young & Stein 1975,
Contr, Biol. & Geol., Milwaukee Pub. Museum, No. 8, 29 p.). Several years of observ-
ing A, fatima populations in the lowland tropical dry forest region (Tosi 1969, Mapa

VOLUME 33, NUMBER 1 59

ecologico de Costa Rica, Trop. Sci. Centr., San José, Costa Rica), Guanacaste Province,
indicate that adults are abundant and active throughout the dry season, especially in
fields with fully leaved shrubs and trees. This note discusses aggregative behavior
of adult A. fatima in Guanacaste.

During the afternoon (1530 h) of 26 January 1977, 12 individuals of A. fatima
were seen fluttering about a low shrub in a field near Playas del Coco. The general
area is about 4 km from the Pacific coast and highly exposed to strong gusty winds
characteristic during the dry season. Most of these butterflies were worn. Eventually
they settled inside the bush for the rest of the day and night. They were scattered
in the leeward side of the bush. Most rested, with wings folded, by hanging from
shaded branches. This behavior was observed again on 20 February 1977 at Playa
Naranjo; six butterflies settled into a shrub at 1615 h remaining there until the fol-
lowing morning. This bush was 400 m from the beach (Gulf of Nicoya) and ex-
posed to strong gusty winds. During the following day, adults fluttered inside the
bush, with occasional settling for short periods (1-6 min). At no time were more
than one or two butterflies resting at once. Both days were sunny, with air tempera-
ture near these bushes being 40-42°C. The air temperature inside them at the same
time of day was 32—34°C. Anartia fatima was active, along with Phoebis ( Pieridae),
in the surrounding habitat, visiting various flowers. These observations took place
during the severe, 4—6 month dry season of this region. The bushes used by A. fatima
were low and dense with branches and leaves. Strong winds coming from the nearby
coast were blocked, as indicated by holding a handkerchief on the lee side of the
bushes.

While it is generally well documented that unpalatable: butterflies exhibit highly
structured communal nocturnal roosting (e.g., Crane 1957, Zoologica 42: 135-146;
Brower & Brower 1964, Zoologica 49: 137-159; Urquart 1960, The Monarch Butter-
fly, Univ. Toronto Press, 361 p.; Owen & Chanter 1969, J. Zool. (London) 157: 345-
374; Young & Thomason 1975, J. Lep. Soc. 29: 243-255), less is known about the
nocturnal behaviors of supposedly palatable species. Other nymphalines, such as
Marpesia berania (Hewitson), Hypolimnas bolina Linnaeus, and Smyrna karwinski
(Geyer), exhibit highly structured nocturnal communal roosting (Barrett & Burns
1951, Butterflies of Australia and New Guinea, Seward, Melbourne, 187 p.; Benson &
Emmel 1973, Ecology 54: 326-335; Muyshondt & Muyshondt 1974, J. Lep. Soc. 28:
924-229) although the adaptive role of this behavior has not been determined. The
unstructured condition of A. fatima aggregates and their low numbers suggests that
such behavior is a sheltering and thermoregulatory response to a dry, windy environ-
ment. Prevailing high temperatures throughout the day, low air humidity, and strong
evening winds induce flocking behavior in A. fatima, causing adults to aggregate in
some bushes. Low, thick bushes offer protection from the sun and evening gusty
winds. On the Pacific slopes of the Cordillera Central in Costa Rica, evening ag-
gregates of another nymphaline, Siproeta stelenes Fabricius, are found in coffee bushes
during the dry season (pers. ob.); these localities are also exposed to strong, gusty
winds and dry conditions.

The presence of worn adults suggests that sites of aggregation are not necessarily
located near eclosion sites; both A. fatima and S. stelenes oviposit singly, and eggs are
distributed over large areas, resulting in low densities of adults eclosing at one spot
(Young & Stein 1975, op. cit., Young & Muyshondt 1974, op. cit.). On Grand Cayman
Island, British West Indies, adults of Anartia jatrophe Linnaeus cling to the leeward
sides of low clumps of the creeping vine Clitoria sp. (Leguminosae) on sunny after-
noons in February (pers. obs. ).

The aggregative behavior of nymphaline butterflies in exposed secondary habitats
near the wind-blown coasts of Caribbean islands and mainland Central America during
the dry season could be related to thermoregulation and physical protection from
strong winds. The dark brown wing and body color of A. fatima and other species
such as S. selenes undoubtedly result in considerable heat gain during afternoon

60 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

hours; the thermoregulatory problems of A. fatima have been discussed (Emmel 1972,
Evolution 26: 96-107). Gradual heat gain on sunny days leads to shade-seeking be-
havior by late afternoon and shady perches also provide nocturnal shelter from winds.
It is known that for some insects living in hot climates, such as desert cicadas, gains
in body heat result in definite periods of movement into shade and subsequent quiet
periods (e.g., Heat & Wilkin 1970, Physiol. Zool. 43: 145-154). Dark butterflies in
lowland tropical dry climates may have similar temperature-response problems.

The above observations and comments suggest that aggregative behavior of A.
fatima is an adaptive response to highly localized climatic or abiotic factors, having
little or nothing to do with biotic factors such as vertebrate predators. A biotic result
of such behavior, however, may be the maintenance of a cohesive adult population
that survives until the following rainy season to oviposit on foodplants as they leaf out.
This idea has been discussed with respect to S. karwinski (Muyshondt & Muyshondt
1974. op. cit.) and it may be generally true for other secondary habitat or pasture-
dwelling tropical nymphalines which pass the dry season in the adult stage.

This fieldwork was funded by N.S.F. Grant GB-33060 and friends of the Museum,
Inc. of the Milwaukee Public Museum.

ALLEN M. Younc, Invertebrate Division, Milwaukee Public Museum, Milwaukee,
Wisconsin 53233.

Journal of the Lepidopterists’ Society
33(1), 1979, 60-64

BISTON BETULARIA, OBLIGATE F. INSULARIA INDISTINGUISHABLE
FROM F. CARBONARIA (GEOMETRIDAE)

It is well known that there is sometimes difficulty in phenotypically recognizing and
scoring f. insularia of Biston betularia. On the one hand it may be confused with
carbonaria (Kettlewell 1973, The Evolution of Melanism, Oxford, London) and
on the other with “typical,” particularly in the Isle of Man (Bailey et al. 1973, En-
tomologist 106: 210-214). To help clarify the matter, various scoring methods have
been devised for insularia, e.g., that of Lees and Creed (1977, Heredity 39: 66-73,
and used by us), where I’ is the lightest and I® the darkest. Neveretheless, difficulties
remain, and Lees and Creed (1977) report a brood, B/574, in which a mating be-
tween two insularia I’/“typical” heterozygotes produced 66 “typical,” 149 T° in-
sularia and 77 carbonaria which were thought to be I*/I®? homozygotes. They also
quote Bowater (1914, J. Genet. 3: 299-315) who crossed a wild carbonaria male with
a “typical” female, and all the resulting progeny were dark insularia, which is con-
sistent with the I*/I* hypothesis.

The present note shows that this cannot always be the explanation since the brood
to be described could not have produced I*/I* homozygotes. On 13 June 1976 a
a female “typical” (Fig. la) was caught in a mercury vapor trap on Hilbre Island near
West Kirby, Wirral, England. She was placed in a tin for the following five nights and
laid a few eggs, but these were infertile. On the night of 19 June 1976, she was put
in a hanging cage with a male insularia (Fig. 1b) (score I’, confirmed by Lees and
Creed, pers. comm.), which had been caught in the mercury vapor trap at Caldy,
Wirral. Mating took place the same evening and the female then laid freely (brood
14672). After she died on 28 June 1976 her body was found to contain a single sper-
matophore only.

61

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64 JoURNAL OF THE LEPIDOPTERISTS SOCIETY

Brood 14672 produced 394 insects, the moths emerging between May and July
1977. There was clear-cut segregation between “typical” (90 6 46; 97 92) and
“dark (100'.6 °¢; 107-99):

Since carbonaria is dominant to insularia, and “typical” is recessive to both, and
because the brood segregated in a 1:1 ratio, “typical” to “dark,” the male parent
must have been insularia/“typical” and the “dark” offspring must all have been
insularia/“typical” heterozygotes.

We found great difficulty in scoring the brood. Some of the insularia males un-
doubtedly had a sparse scattering of white scales and rather light hindwings (Fig. 2a),
but they lacked the white scales on the thorax or abdomen characteristic of insularia.
Others appeared to us to be quite indistinguishable from carbonaria and we cannot
agree that the two can easily be separated, “. . . since in carbonaria, when the white
scales are present on the forewing, they are distributed to form a thin, unbroken trans-
verse line across the middle of the forewing” (Steward 1977, Ecol. Entomol. 2: 231-
243). The insularia females in brood 14672 (Fig. 2b) present even more of a
problem, because they are normally darker than the males and we should have scored
nearly all of them as carbonaria (Figs. 3b and 4b).

We appreciate that in expert hands specimens may be scored more competently,
but had the “dark” males of brood 14672 been “trap caught” (Fig. 3a), we (and we
think others) would have scored many of them (see Fig. 4a) as “carbonaria,’ and
Creed (pers. comm.) agrees with this view.

Kettlewell (1973, The Evolution of Melanism, Oxford, London) thinks that those
insularia which are difficult to distinguish from carbonaria are in a minority,
as are the pale insularia misclassified as dark “typical.” He further thinks that be-
cause of the small numbers in these two categories, the frequency figures for “typical”
and insularia are largely unaffected. However, it is difficult to be sure of this since
we do not know how often misscoring takes place, and different observers in the same
locality do, in fact, obtain very different results when recording the frequency of the
three forms (see Kettlewell, 1973, op. cit., table 9: 2).

Creed (pers. comm.) thinks that carbonaria as scored in the wild is perhaps as
heterogeneous as insularia. We agree with this, and feel that the possibility of mis-
scoring of males should be seriously considered when unusual frequencies of car-
bonaria and insularia are reported. (The misscoring of females is much less im-
portant as they rarely come to mercury vapor light and never to an assembly trap. )

Sib matings and backcrosses using brood 14672 were set up and their results are
given in Table 1. Moths which were obligatory insularia/“typical” heterozygotes
again were almost always indistinguishable from wild-caught carbonaria, as were
insularia/carbonaria heterozygotes. Furthermore, there was no detectable differ-
ence between any of these “dark” insects and those in brood 15088, which must have
contained insularia homozygotes. An additional mating was set up from this stock
in 1978 using the insularia female (I*) from brood 15090. These results will be
reported later. |

We are grateful to the late Dr. E. R. Creed and to Dr. D. R. Lees for scoring a
sample of our brood and for their helpful comments, and to the Nuffield Foundation,
The Royal Society and the Science Research Council for continued support.

Note added in proof: Brood 15490 using this female I® mated to a male “typical” is
segregating (so far) cleanly I* and “typical” with no carbonaria.

Sin Cyr A. CLArke, Hon. Research Fellow, Department of Genetics, University
of Liverpool, England.

Journal of the Lepidopterists’ Society
33(1), 1979, 65-67

BROTHER APOLINAR MARIA (1867-1949) AND HIS CONTRIBUTIONS
TO COLOMBIAN LEPIDOPTEROLOGY

Brother Apolinar Maria (né Nicholas Seiler) is an all but forgotten name in the
annals of South American Lepidopterology. Nevertheless, he built up the largest
collection of butterflies in Colombia. He published several papers on Lepidoptera
and described a number of new taxa. He sent countless Colombian specimens to sev-
eral institutions and private collections in the Americas and Europe. This note gives
an account of his life and deeds, and the butterfly taxa he described.

Brother Apolinar Maria was born on November 5th, 1867 in Sarreguemines, Alsace,
France. He lived for some time in Reims, and possibly it was here that he joined
the religious Brotherhood of La Salle. He was sent by his superior to Colombia, where
he arrived in 1904 on the S.S. “Leon XIII.”

A few years later (in 1912) he founded the Natural History Society of the La Salle
Institute in Bogoté, with the enthusiastic help of some local amateur naturalists,
several of whom had been his own pupils. The Society published the first number
of its periodical in 1913, and it went on uninterruptedly until 1931, when it stopped
publication after 110 numbers.

In its pages, Brother Apolinar, his colleagues, pupils and friends published numerous
observations on Colombian natural history. Among short notes and some longer
papers, Brother Apolinar made 75 contributions on Colombian butterflies.’

Through numerous correspondents in many regions of Colombia, Brother Apolinar
received large numbers of natural history objects, which were used by him to enrich
the holdings of the Natural History Museum. This museum had become by far the
most important natural history institution of Colombia at that time. By 1930, the
Lepidoptera collection included 17,235 specimens.

Many butterfly specimens also were sent to friends in America and Europe. Among
his many correspondents were C. Oberthiir, A. H. Fassl, W. Schaus, H. G. Dyar and
R. F. d’ Almeida. Later, Apolinar was elected an honorary life member of the Colom-
bian Academy of Exact, Physical and Natural Sciences.

In 1948 he received the severest blow in his long scientific career. On April 9th
and 10th rioting and plundering parties set fire to, and destroyed, a great number of
public and private buildings in downtown Bogota. The library and the collections of
the La Salle Natural History Museum were irretrievably lost (cf. Dugand 1948,
Caldasia 5(22): 223; LeMoult 1948, Misc. Entomol. 45: 93; Remington & Reming-
ton 1948, Lepid. News 2: 61; Ruiz 1950, Rev. Acad. Colomb. Cienc. Exact. Fis.
Nat. 7(28): 433; and Daniel 1950, Bol. Soc. Cienc. Nat. Caldas 51: 74). Apolinar
~ died on December 24th, 1949.

Between 1914 and 1942 Brother Apolinar described 42 butterfly taxa. Several of
those taxa have never been recorded in the Zoological Record (nor by Beattie 1976,
Rhopalocera Directory). In spite of the rather naive form of his taxonomic descrip-
tions, they are nevertheless valid, and his names are available in most cases, so that
they should be accepted as representing valid taxa. Unfortunately, his descriptions are
usually quite vague. In most instances no figures were provided, and all of his types
were lost in the 1948 fire.

List oF BUTTERFLY TAXA DESCRIBED BY BROTHER APOLINAR MARIA

BLS = Boletin de la Sociedad de Ciencias Naturales del Instituto de La Salle;
BSC = Boletin de la Sociedad Colombiana de Ciencias Naturales; RSC = Revista de
la Sociedad Colombiana de Ciencias Naturales; RAC = Revista de la Academia Co-
lombiana de Ciencias Exxactas, Fisicas y Naturales.

1A bibliography of Apolinar’s publications on Colombian butterflies is available from the author
upon request.

66 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

PAPILIONIDAE

Papilio isidorus var. leleargei; BLS, 1915, 3(9): 142 (Susumuco, Colombia) [Holo-
type; sex not stated] [“leleargei” is a typographical error for “lelargei”].

Papilio oberthuri; BLS, 1916, 4(27): 32 (Quebrada Negra, El] Baldio, Colombia)
[ Holoytpe; sex not stated].

Papilio euryleon abadiae; BLS, 1916, 4(31): 95 (Muzo, Colombia) [Holotype; sex
not stated].

Papilio rhodostictus nymphius “orientalis’; BSC, 1924, 13(76): 108 (“Llanos,”
Villavicencio, Medina, etc., eastern Colombia) [14 ¢ ¢ syntypes].

Papilio rhodostictus nymphius “occidentalis”; BSC, 1924, 13(76): 108 (Mauzo,
Colombia) [5 ¢ 2,1 9, syntypes].

PIERIDAE

Colias dimera mariae; BLS, 1914, 2(1): 17 (Serrania de Monserrate, Paramo de
Choachi, Colombia, 2900 m, 2.1.14) [Holotype; sex not stated].

Colias dimera var. fassli; BLS, 1914, 2(1): 17 (Paramo de Choachi, Colombia) [sev-
eral ¢ and @ syntypes; figured in RAC, 1942, 5(17): pl. [1], figs. 9-10]

Nathalis plauta var. nigrescens; BLS, 1914, 2(2): 45 (San Cristébal, Colombia,
31.xii.13) [Holotype ¢ ].

Nathalis plauta var. xanthoptera; BLS, 1914, 2(10): 304 (neighborhood of Bogota,
Colombia) [several 9 syntypes].

Perrhybris pandora ab. “teresa”; BSC, 1926, 15(84): 15 (Villavicencio, Colombia )
Holotype 92; figured as P. p. var. teresa in RAC, 1942, 5(17): pl. [1], fig. 4.

Perrhybris lypera ab. “mariae”; RSC, 1930, 4(105): 59 (Muzo, Colombia) [two
syntypes; figured in RAC, 1942, 5(17): pl. [1], fig. 3].

Pereute leucodrosime ab. “mariae”; BSC, 1926, 15(84): 17 (Villavicencio, Colom-
bia) [Holotype 9; figured as P. l. var. mariae in RAC, 1942, 5(17): pl. [1], fig 5].

Catasticta uricoecheae bouwvieri; BSC, 1926, 15(84): 24 (Chocd, Colombia) [Holo-
type; sex not stated; figured as C. bouvieri in RAC, 1942, 5(17): pl. [1], fig. 8].

Daptonoura calymnia ab. “leucoptera’; BSC, 1926, 15(85): 43 (Las Mesitas, Fusa-
gasuga, Colombia) [two syntypes; sex not stated].

Daptonoura polhymnia flavopunctata; BSC, 1926, 15(85): 44 (Muzo, Colombia)
[4 syntypes; sex not stated].

Teria [sic] mexicana henricii; BSC, 1926, 15(85): 46 (Sonsén, Colombia) [3 ¢ é,
2 2 2, syntypes].

Teria [sic] gaugamela alba; BSC, 1926, 15(85): 47 (Medellin, Colombia) [Holo-
type 2].

Dismorphia acutipennis llerasi; BSC, 1926, 15(86): 96 (Llanos de Guaicaramo, Co-
lombia) [Holotype; sex not stated; figured in RAC, 1942, 5(17): pl. [1], fig. 6].

Dismorphia arcadia ab. “melanoptera’”; BSC, 1926, 15(86): 99 ( Villavicencio; Pen-
silvania, Caldas, Colombia) [1 6,1 9, syntypes].

Dismorphia demeter confluens; BSC, 1926, 15(86): 100 (Vergara or Choachi, Co-
lombia) [Holotype; sex not stated].

NYMPHALIDAE
SATYRINAE

Idioneura erebioides £. intermedia; BLS, 1914, 2(2): 46 (no locality) [several syn-
types; sex not stated].

Lymanopoda samius var. nigripunctata; BLS, 1914, 2(2): 47 (no locality) [Holo-
type; sex not stated].

Lymanopoda samius var. confluens; BLS, 1914, 2(2): 48 (no locality) [several syn-
types; sex not stated].

Pedaliodes nebris albipunctata; BLS, 1914, 2(3): 76 (Paramo de Choachi, Colom-
bia, 3100 m) [Holotype; sex not stated].

VoLUME 33, NUMBER 1 67

Pedaliodes nebris athymi; BLS, 1914, 2(3): 76 (Paéramo de Choachi, Colombia,
3100 m) [15 syntypes; sex not stated].

Pedaliodes nebris conchae; BLS, 1914, 2(3): 77 (Paéramo de Choachi, Colombia,
3100 m) [10 syntypes; sex not stated].

Pedaliodes nebris var. abadiae; BLS, 1914, 2(3): 77 (Paramo de Choachi, Colombia,
3100 m) [3 syntypes; sex not stated].

Pedaliodes nebris var. pauli; BLS, 1914, 2(3): 77 (Paramo de Choachi, Colombia,
3100 m) [10 syntypes; sex not stated].

Pedaliodes nebris var. tripunctata; BLS, 1914, 2(3): 77 (Paramo de Choachi, Co-
lombia, 3100 m) [several syntypes; sex not stated].

Pedaliodes nebris var. modesta; BLS, 1914, 2(3): 77 (Paramo de Choachi, Colom-
bia, 3100 m) [Holotype; sex not stated].

Pedaliodes nebris estanislaoi; BLS, 1914, 2(3): 78 (Paramo de Choachi, Colombia,
3100 m). [Holotype; sex not stated].

Zapatoca; BSC, 1924, 13(76): 84. Unjustified emendation of Sabatoga Staudinger,
1897. Invalid.

Zapatoca [sic] viventieni; BSC, 1924, 13(76): 84 (Guasca, Colombia) [Holotype;
sex not stated].

MoORPHINAE

Morpho amathonte var. nigromarginata; RAC, 1942, 5(17): pl. [1], fig. 2 (no local-
ity) [Holotype ¢ ].

ITHOMIINAE

Mechanitis egaensis septentrionalis; BSC, 1928, 17(98): 164, 180 (Valle de Tensa,
Garagoa, Boyaca, Colombia [Holotype; sex not stated].

HELICONIINAE

Heliconius ismenius abadiae; BSC, 1926, 15(87): 125 (no locality) [3 syntypes; sex
not stated].

Heliconius cydno hermogenes ab. “xanthosticta”; BSC, 1926, 15(88): 153 (Valle del
Cauca, Colombia ) [11 syntypes; sex not stated].

Heliconius cydno hermogenes ab. “leucosticta”; BSC, 1926, 15(88): 153 (Valle del
Cauca, Colombia ) [19 syntypes; sex not stated].

Heliconius cydno dolores; BSC, 1926, 15(88): 155 (Tolima, Colombia) [Holotype;
sex not stated].

Heliconius aristiona colombiana; BSC, 1927, 16(92): 117 (Susumuco, Colombia )
[ Holotype; sex not stated].

Heliconius mixta; BSC, 1927, 16(92): 119 (Llanos de Medina, Guaicaramo, Colom-
bia) [Holotype; sex not stated].

NYMPHALINAE

Callicore astala var. coeruleomarginata; BSC, 1928, 17(98): 164, 183 (Rio Putu-
mayo, Colombia) [Holotype; sex not stated].

GERARDO LAMAS, Museo de Historia Natural “Javier Prado,’ Universidad Nacional
Mayor de San Marcos, Apartado, 1109, Lima-100, Pert.

Journal of the Lepidopterists’ Society
33(1), 1979, 68-69

WEATHER AND THE REGULATION OF HYPOTHYRIS EUCLEA
(NYMPHALIDAE ): POPULATIONS IN NORTHEASTERN COSTA RICA

Brown and Neto (1976, Biotropica 8: 136-141) found that populations of some
ithomiine butterflies (Hypothyris and Mechanitis) in Brazil are controlled largely by
parasitism of eggs and larvae during the wet season. Presumably periods of increased
daily precipitation provide increased opportunities for successful parasitic attacks.
Characteristically, local adult populations of Mechanitis and Hypothyris exhibit large
fluctuations in numbers throughout the year (Brown & Neto, op. cit.; pers. obs. ).

Populations of Hypothyris euclea leucania (Bates) exhibit annual periods of sud-
den, rapid growth of adult populations in northeastern Costa Rica (Young 1977, Pan-
Pacif. Entomol. 53: 104-113). It was presumed in that study that the frequency of
mating and oviposition and the survival of eggs and larvae were increased during a
period of dryness preceding the time of increased population abundance. The pur-
pose of this note is to present further data on H. euclea that support my earlier pre-
diction that greatly increased adult numbers, in northeastern Costa Rica, follow dry
periods.

At times each year (1971-77), at Finca La Tirimbina, La Virgen, Heredia Prov-
ince (220 m elev.), very high density concentrations of adult H. euclea occur along
forest trails and clearing. At these times, many individuais of the larval foodplant,
Solanum rugosum Dund. (Solanaceae), are heavily defoliated by H. euclea larvae
feeding a few weeks earlier. Few larvae are present during weeks of high adult
abundance. The deposition of many large egg masses on S. rugosum results in many
larvae and defoliation ( Young, op. cit.). Walking along a forest trail flushes out many
resting butterflies; from one to 30 adults may be flushed from a five-meter section
of trail on a sunny day. Most of these are fresh. At other times of the year, as few as
one or two adults occur in about 100 meters of trail. Sometimes adults are conspicu-
ously absent. Adults feed on dead insects and fresh bird droppings. Either adult
dispersal or mortality is high over short periods (one week) since, for example, from
AO fresh adults marked during a 20-minute period one day (15 August 1977) at

580
540 |
500
460
420
380
ma
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E 300+
= 260+
5
220
180}
140"
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60+
20
1 5 10 15 20 25 30 1 5 10 15 20
July August

SUCCESSIVE DAYS

Fic. 1. Daily rainfall levels for July and August 1977 at Finca La Tirimbina, La
Virgen de Sarapiqui. Note the two-week dry period in the latter part of July. Data
courtesy Dr. J. Robert Hunter,

VoLUME 33, NUMBER 1 69

one spot, only three were recaptured two days later and none a week later. Two-
thirds of these butterflies were males. A few weeks later, adult numbers had de-
clined considerably at five different spots where abundances were very high for at
least three weeks. Brown and Neto (op. cit.) found that populations of H. euclea
and H. daeta diminish rapidly in size as a result of high dispersal and high longevity
of adults. If July-August 1977 is used as an example, August was characterized by
heavy daily downpours, both day and night, but this rainy period was preceded by
a dry period during the last two weeks of July ( Fig. 1).

The life cycle of H. euclea at this locality takes about 22 days (Young, op. cit.).
Thus frequent deposition of large egg masses at this locality and high survival of eggs
and larvae result in a large wave of fresh adults about three weeks later. For ex-
ample, egg masses deposited during the dry period in July 1977 produced the large
adult population present in August. Mortality factors operative on egg rafts and gre-
garious young larvae may be drastically reduced in frequency and intensity during
periods of dry weather (Young, op. cit.). Although the proximal causes of this ap-
parent mortality are unknown, their activity correlates well with wet periods. A broad
range of invertebrate predators and pathogenic fungi are very likely involved in the
regulation of H. euclea populations. Gilbert (1969, Some aspects of the ecology and
community structure of ithomiid butterflies in Costa Rica, Organization for Tropical
Studies, mimeo report) found ants and wasps to be predators of H. euclea larvae.
Gilbert also suggests that egg mortality from leaf-patrolling predators is operative in
H. euclea populations. Waves of pupal and adult flour beetles (Tribolium spp.) fol-
low periods of slackened predation (Mertz 1969, Ecol. Mongr. 39: 1-31. Dry
weather may also enhance mating and oviposition (Young, op. cit.), thereby “stacking
the deck” even further for a large cohort of adults to appear.

These observations suggest that tropical butterfly populations subject to control
or regulation by biotic agents may be, in fact, regulated only to the extent to which
daily rainfall patterns influence the activity of these agents.

ALLEN M. Younc, Invertebrate Division, Milwaukee Public Museum, Milwaukee,
Wisconsin 53233.

Journal of the Lepidopterists’ Society
33(1), 1979, 69-71

BOOK REVIEW

PENNINGTON’S BUTTERFLIES OF SOUTHERN AFRICA, edited by C. G. C. Dickson with D.
M. Kroon, 1978. Ad. Donker, Johannesburg and London. 671 pp., including frontis-
piece, 198 plates, 1 text figure and 1 map. Price: R49.00 (approximately $80.00
WS.)

The manuscript for this book was begun many years ago by K. M. Pennington, but
“KMP” never lived to see its appearance. It is a testimony to the heartfelt admiration
that his fellow collectors felt toward Pennington that they were determined that
KMP’s life work would not go unpublished. For the next three years Mr. Dickson
and Dr. Kroon revised, updated and added to the manuscript to make it ready for
publication. They have done an admirable job and produced in Pennington’s honor
a truly outstanding book, one that is not duplicated by other regional treatments of
a comparably sized fauna.

The 781 species of butterflies found in southern Africa (including South Africa,
Southwest Africa and much of adjacent Rhodesia and Mozambique) are treated in
the text on pages 33-201. This is a fauna comparable to the rhopaloceran fauna oc-

70 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

curring in the United States and Canada. The treatment includes information on
where, when and in what habitats one may find any South African butterfly. Original
descriptions are cited meticulously and accurately for all species and many of their
synonyms—I only wish in a couple of instances that the editors had used the senior
title for works (such as Die GrossSchmetterlinge der Erde instead of the later transla-
tion The Macrolepidoptera of the World by Seitz, and De Uitlandsche Kapellen .. .
rather than Papillons Exotiques . .. for Cramer), though in both of these instances
footnotes show that the editors accepted the correctness of the original titles but were
using those customarily employed in past English language publications.

The plates are excellent, very informative and of the highest technical quality.
Using them to identify a South African butterfly is a joy compared with the aggrava-
tions the reader is faced with in some recent major books on other faunas. The illus-
trations in this book, along with those in the equally wonderful Butterflies of Japan
(T. Fujioka, 1975) are the best advertisements for using quality colored photographic
plates that I can imagine. There is no possibility of confusing similar species (as-
suming that superficial characters will distinguish them) by use of these plates and
the accompanying text. I might have preferred to see the backgrounds a bit paler
blue, but this is just a personal complaint; in no way deo the backgrounds detract
from the plates’ usefulness.

Dr. Kroon’s list of the foodplants of South African butterflies (pages 604-643) is
a fine and useful compilation of what is known about the life cycles of the region’s
Rhopalocera. The introductory portion (pages 22-30) by Mr. Dickson show that the
South African collectors are far more interested in life histories than are we in this
country, and one suspects that they are near the top of the world in awareness of this
aspect of lepidopterology.

In any event, the text and plates combined make the identification of any South
African butterfly at least possible, something that could not have been said before
the publication of the present book.

But no book is perfect, as the editors would be the first to admit, and this one is no
exception. Very few typographical errors or mistaken facts have crept into these
pages, but the editors might profitably have looked to this side of the Atlantic for
some higher classificatory schemes. Munroe’s papilionid, Klots’ pierid and my satyrid
higher classifications are not mentioned, much less used; some relationships could
have been elucidated had they been. Fox demonstrated rather well that Sallya
(=Crenis of authors) is really only a glorified Eunica, at least the boisduvali com-
plex is, but since this work was not cited, the conclusions were apparently not seen.
More surprising, though, is the use of Meneris instead of Aeropetes for the spectacular
satyrid A. tulbaghia (Linné). That generic synonymy was pointed out years ago by
Hemming who resurrected Billberg’s 1820 work from obscurity, thus correctly re-
placing some well-known generic names with Billbergian ones.

The plates, excellent as they are, have not totally escaped the inevitable “glitches.”
Plate 76 is especially affected since wet and dry season forms are uniformly mixed and
Figure 146.ix is of a dry season individual, but this fact is not noted.

Most of the specimens illustrated are in the collection of the Transvaal Museum in
Pretoria, but a few are taken from material in other museum and private collections.
All sources are cited in the captions. The figure numbers are cross-referenced to the
running catalog of the species in the text. This makes finding the text material re-
ferred to by the plates very easy, and it also gives an idea of how many of each
species are figured. For example, there are 15 specimens of Charaxes zoolina zoolina
(species 146 in the text) figured, and they are marked by figure numbers 146i to
146xv on the plates. On the occasions that additional specimens are figured out of
sequence, this fact is noted where the specimen might have been expected, so the
gynandromorph of Colotis ione (species 606) is figured on Plate 196 as figure 606xxv,
but it is cross-referenced on Plate 150 where it logically would have been placed.
Parenthetically, the symbol used in this book for a gynandromorph (¥% ) is that tra-

VoLUME 33, NUMBER 1 7a

ditionally used for a worker caste of a social insect, rather than the usual ¢ em-
ployed for these oddities. The fact that so many examples are figured in variable
species enables one to see at a glance the expected variation—a further aid in iden-
tification.

The first 38 plates are taken from the late G. C. Clark’s incomparable life history
studies, encompassing the cycles of 37 species (there are two of Borbo fatuellus from
different populations) of Hesperiidae. They are of the same quality and in the same
style as his earlier plates on the Papilionoidea of South Africa, and Clark, too, has
left an impressive legacy to future lepidopterists.

The shortcomings pointed to here in no way diminish the value of this book. It
is a major accomplishment, and one that has quickly changed working with South
African butterflies from one of the least, to one of the most, possible tasks. Everyone
connected with this book—Mr. Dickson, Dr. Kroon, the Trustees of the Ken Penning-
ton’s Butterflies of Southern Africa Trust and the printer—are to be congratulated on
producing a magnificent major work. I think that K. M. Pennington would be proud
of it!

Lee D. Mitter, Allyn Museum of Entomology, 3701 Bay Shore Rd., Sarasota,
Florida 33580.

Journal of the Lepidopterists’ Society
33(1), 1979, 72-75

OBITUARY

tH SS ae

CHRISTOPHER HENNE (1905-1977 )

The world of lepidopterology lost a great friend and lepidopterist when Christopher
Henne, a long time member of the Lepidopterists’ Society, passed away on October
o, 1977,

Henne was born in Denver, Colorado, on July 20, 1905, and at 18 months of age
he and his parents moved to California. He grew up in Pasadena where he attended
the Polytechnic Elementary School and the Pasadena Military Academy. In 1920, he
went to Europe with his mother and spent two years in an English preparatory school
in Switzerland. At the age of 17, he returned to the United States and finished his
schooling at the Asheville School in Asheville, North Carolina. He then returned to
southern California where he was to reside the rest of his life. .

He was self-taught as far as his background in entomology was concerned, for he
had never even had a course in general biology. He worked in a variety of entomo-
logically related positions, including display preparation and teaching at the Los An-
geles County Museum, later followed by the establishment of his own business, Henne
Biological Supply, in which he practiced plastic embedding of insects for educational
institutions.

In 1961, Chris and his wife Dorothy moved to a new home at the edge of the Mojave
Desert in Pearblossom, California, where they retired to full-time collecting. During
the next 16 years, his collection was to more than quadruple in size.

VoLUME 33, NUMBER 1 7h

Henne’s collecting started at age four when the family gardener gave him a Cecropia
Moth for Christmas. This stimulated a lifelong interest and specialization in moths
and butterflies. He soon drove his parents to distraction by his habit of removing
the bathroom window screen at night and using the bathroom as a giant light trap for
the local moths. As a teenager in Switzerland, he met his first professional lepidop-
terist, an Englishman, who took him on collecting trips.

After leaving Asheville and returning to Pasadena, Henne was to become one of
the most diligent and thorough explorers of the southern California deserts and moun-
tains, collecting in company with most of the noted California lepidopterists of the
next five decades. Among his companions during these years of many exciting lepidop-
terological discoveries were John Adams Comstock, Charles M. Dammers, Jean Gun-
der, Charles Ingham, Lloyd Martin, C. N. Rudkin, and Munroe Walton. Collecting
companions and visitors to his home in recent years included numerous members of
the Pacific Slope Section of the Lepidopterists’ Society. One could rarely drop in on
the Hennes in their Pearblossom home without running into another lepidopterist
enjoying their hospitality in the form of “Hennes-on-the-Rocks” (a rum and limeade
drink) or “Entomological Goop” (a Noodles Romanoff concoction of shrimp, mush-
rooms and pimentoes ).

Henne’s first love was desert collecting and it was in the far reaches of the Colorado
and Mojave deserts that he made his most interesting discoveries in Lepiodptera.
He was one of the first collectors to investigate the Providence and New York Moun-
tains, and he continued to make field trips to these desert ranges up to the time of his
death. In the region around Pearblossom, he studied the biology of numerous butter-
fly and moth species, recording an immense quantity of information about their food-
plants and life histories.

In collaboration with his good friend John Comstock, he worked out many hitherto
unknown life histories. Henne collected this material, obtaining ova, larvae, and
pupae, and sent it to Comstock who then prepared paintings and descriptions for joint
publication. With Comstock’s death in 1970, this partnership ceased, but Henne con-
tinued to amass data which will be valuable to current and future workers studying
the California butterfly and moth faunas. Much of this information is recorded on
the very detailed specimen labels on pinned adults in his collection.

He was particularly interested in diurnal moths, and this is reflected in the rich
representation of diurnal species of Pyralidae, Ctenuchidae, Sesiidae, and Noctuidae in
his collection. In this last family, he took a special interest in the genus Annaphila, of
which he collected or reared large series of most of the California species. The plume
moths (Pterophoridae) were another favorite of his, as evidenced by hundreds of
specimens in his collection, many of which were reared.

One of his greatest entomological finds was the rediscovery of the day-flying
sphingid, Euproserpinus euterpe Henry Edwards, formerly known from only a hand-
ful of specimens collected some 90 years ago. He collected a series of specimens while
establishing the habitat and foodplant of this rare species.

He named two species, one subspecies (with Comstock), and one form of Lepidop-
tera, as follows: Callophrys comstocki Henne (Lycaenidae); Copicucullia mcdun-
noughi Henne (Noctuidae); Philotes enoptes dammersi Comstock & Henne ( Lycaeni-
idae); and Leptotes marina form “burdicki’ Henne (Lycaenidae). Insects named
after him are: Euphydryas chalcedona quino transition form “hennei” Gunder (Nym-
phalidae); Speyeria coronis hennei (Gunder) (Nymphalidae); Penstemonia hennei
Engelhardt (Sesiidae); Abagrotis hennei Buckett (Noctuidae); Annaphila (Proanna-
phila) hennei Rindge & Smith (Noctuidae); Racheospila hennei Sperry (Geometri-
dae) (NOW Nemoria obliqua hennei (Sperry); Eucosma hennei Clarke (Tortricidae);
and Cophura hennei Wilcox (Diptera: Asilidae).

The Henne collection was purchased in December 1977, by the Los Angeles County
Museum of Natural History Foundation. It consists of 36,896 specimens, of which
approximately 88% are moths and 12% are butterflies. In the moths, the families

74 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Noctuidae and Geometridae are best represented, along with relatively large num-
bers of Pterophoridae, diurnal Pyralidae, Ctenuchidae, and Sesiidae. In the butter-
flies, the genera Apodemia, Philotes s.1., and Euphydryas are particularly well repre-
sented. Each specimen is receiving an accession label and is being incorporated into
the general collection of the museum, where all will be available for examination and
study.

ee to Lepidoptera, Henne loved animals, especially dogs. Two of his pet dogs
were orphans he found on collecting trips. He also had a keen interest in auto-
mobiles, and frequented car shows and races throughout southern California. Pursuing
two hobbies simultaneously, his early trips over primitive desert roads to the famous
Bonanza King Mine area of the Providence Mountains were made in a prized 1934
Auburn Speedster.

His inventive mind developed a number of novel techniques for rearing in his Pear-
blossom laboratory. One of the spin-offs was the development of the Henne Clothes-
Moth Trap. He also developed techniques in the field of plastic embedding which
retained the natural color and shape of the insects used.

In everything he did, whether rearing larvae or spreading adult specimens, Chris
Henne was a perfectionist. His collection of immaculately curated material, with in-
credibly detailed data, is as aesthetically pleasing as it is scientifically valuable. Any-
one who sees this collection will immediately appreciate the copious time and effort
put into it by one of California’s most prolific and gifted lepidopterists. He will be
greatly missed by all who had the privilege and pleasure of knowing him.

JoHN F. EMMEL, 41783 El Camino Drive, Hemet, California 92343.
THomas C. EMMEL, Department of Zoology, University of Florida, Gainesville,
Florida 32611.

PUBLICATIONS OF CHRISTOPHER HENNE

1. Comstock, J. A. & C. Henne. 1933. A new lycaenid from southern California.
Bull. So. Calif. Acad. Sci. 32: 23-26.

2. Henne, C. 1935. A new form of Leptotes marina (Lepid.: Nymphalidae)
(sic). Entomol. News 46: 100-101.

. 1940. Two new species of Lepidoptera from California. Bull. So.
Calif. Acad. Sci. 39: 71-74.

4. Comstock, J. A. & C. Henne. 1940. Notes on the early stages of Nemoria
pistaciaria Pack. Bull. So. Calif. Acad. Sci. 39: 78-80.

Bp 1940. A comparison of the larva of Arachnis picta insularis and
Arachnis picta maia. Bull. So. Calif. Acad. Sci. 39: 189-190.

6. 1940. Notes on the early stages of Xanthothrix ranunculi. Bull. So.
Calif. Acad. Sci. 39: 198-199.

if 1941. The larva and pupa of Trichoclea edwardsi Sm. Bull. So. Calif.
Acad, Sci. 40: 165-166.

8. 1942. Notes on the life history of Tolype glenwoodii Barnes. Bull.

So. Calif. Acad. Sci. 41: 86-90.

. 1942. The early stages of Arctonotus lucidus Bdy. (Lepidoptera).

Bull. So. Calif. Acad. Sci. 41: 167-171.

. 1943. The larva of Copicucullia basipuncta B. & McD. (Lepidoptera).

Bull. So. Calif. Acad. Sci. 42: 45-46.

1943 (1944). Mature larva of Graptolitha longior Sm. Bull. So.
Calif. Acad. Sci. 42: 132.

12. Henne, C. 1948. More notes on California winter Lepidoptera. Lepid. News
2: 42.

. 1957. Charles Henry Ingham (1904-1957) (obituary). Lepid. News
11: 169-170.

VOLUME 33, NUMBER 1 75

14.

15.
16.

Ne

18.
19.
20.
21.

22.

23.
24.

25.

Comstock, J. A., C. Henne & F. Sala. 1957 (1958). The habits and life his-
tories of Cochisea sinuaria and Cochisea sonomensis. Bull. So. Calif. Acad. Sci.
56: 169-177.

Comstock, J. A. & C. Henne. 1964. Studies in life histories of North American
Lepidoptera. California Annaphilas. J. Res. Lepid. 3: 175-191.

1965. Notes on the life history of Philotes enoptes dammersi. Bull.
So. Calif. Acad. Sci. 64: 153-156.

Hogue, C., F. Sala, N. McFarland & C. Henne. 1965 (1966). Systematics and
life history of Saturnia (Calostaturnia) albofasciata in California (Saturni-
idae). J. Res. Lepid. 4: 173-184.

Comstock, J. A. & C. Henne. 1966. Studies in life histories of North American
Lepidoptera. California Annaphilas II. J. Res. Lepid. 5: 15-26.

. 1966. The larva and pupa of Orthosia hibisci quinquefasciata (Noc-

tuidae). J. Lepid. Soc. 20: 213-215.

1967. Notes on the life history of Philotes rita elvirae (Lepidoptera;

Lycaenidae ). Bull. Soc. Calif. Acad. Sci. 66: 99-102.

. 1967. Early stages of Sphinx sequoiae engelhardti (Sphingidae). J.
Lepid. Soc. 21: 27-31.

Henne, C. 1967 (1969). Field investigations preliminary to life history
studies on the lithosina-miona-casta complex of the genus Annaphila (Noctui-
dae). J. Res. Lepid. 6: 249-256.

Comstock, J. A. & C. Henne. 1967 (1969). Studies in life histories of North
American Lepidoptera. California Annaphila III. J. Res. Lepid. 6: 257-262.

1967 (1969). Early stages of Lycomorpha regulus Grinnell, with notes

on the imago (Lepidoptera: Amatidae). J. Res. Lepid. 6: 275-280.

1969. Life history notes on Lithophana subtilis (Noctuidae). J.

Lepid. Soc. 23: 15-18.

EDITORIAL STAFF OF THE JOURNAL
AUSTIN P. PLATT, Editor
Department of Biological Sciences
University of Maryland Baltimore County, 5401 Wilkens Avenue
Catonsville, Maryland 21228 U.S.A.

FRANCES S. CHEW, Managing Editor

Department of Biology
Tufts University
Medford, Massachusetts 02155 USA

DouGLas C. FERGUSON, Associate Editor THEODORE D. SARGENT, Associate Editor

NOTICE TO CONTRIBUTORS

Contributions to the Journal may deal with any aspect of the collection and study of
Lepidoptera. Contributors should prepare manuscripts according to the following in-
structions.

Abstract: A brief abstract should precede the text of all articles.

Text: Manuscripts should be submitted in triplicate, and must be typewritten,
entirely double-spaced, employing wide margins, on one side only of white, 8% x 11
inch paper. Titles should be explicit and descriptive of the article’s content,including
the family name of the subject, but must be kept as short as possible. The first mention
of a plant or animal in the text should include the full scientific name, with authors
of zoological names. Insect measurements should be given in metric units; times
should be given in terms of the 24-hour clock (e.g. 0930, not 9:30 AM). Underline only
where italics are intended. References to footnotes should be numbered consecutively,
and the footnotes typed on a separate sheet.

Literature Cited: References in the text of articles should be given as, Sheppard
(1959) or (Sheppard 1959, 1961la, 1961b) and all must be listed alphabetically under
the heading LITERATURE CITED, in the following format:

SHEPPARD, .P. M. 1959. Natural selection and heredity. 2nd. ed. Hutchinson, London.
209 p.

196la. Some contributions to population genetics resulting from the study of
the Lepidoptera. Adv. Genet. 10: 165-216.

In the case of general notes, references should be given in the text as, Sheppard (1961,
Adv. Genet. 10: 165-216) of (Sheppard 1961, Sym. R. Entomol. Soc. London 1: 23-
30).

Illustrations: All photographs and drawings should be mounted on stiff, white
backing, arranged in the desired format, allowing (with particular regard to lettering)
for reduction to their final width (usually 4% inches). Illustrations larger than 84% x 11
inches are not acceptable and should be reduced photographically to that size or small-
er. The author's name, figure numbers as cited in the text, and an indication of the
article’s title should be printed on the back of each mounted plate. Figures, both line
drawings and halftones (photographs), should be numbered consecutively in Arabic
numerals. The term “plate” should not be employed. Figure legends must be type-
written, double-spaced, on a separate sheet (not attached to the illustrations), headed
EXPLANATION OF FIGURES, with a separate paragraph devoted to each page of illus-
trations.

Tables: Tables should be numbered consecutively in Arabic numerals. Headings
for tables should not be capitalized. Tabular material should be kept to a minimum and
must be typed on separate sheets, and placed following the main text, with the ap-
proximate desired position indicated in the text. Vertical rules should be avoided.

Proofs: The edited manuscript and galley proofs will be mailed to the author for
correction of printer's errors. Excessive author's changes at this time will be charged
to authors at the rate of 75* per line. A purchase order for reprints will accompany the
proofs.

Correspondence: Address all matters relating to the Journal to the editor. Short
manuscripts such as new state records, current events, and notices should be sent to
the editor of the News: Jo Brewer, 257 Common Street, Dedham, Massachusetts 02026
U.S.A.

.

PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044 U.S.A.

CONTENTS

PAITITIA NEGLECTA, GEN. N., SP. N. FROM PERU (NYMPHALIDAE:
ITHOMIINAE). Gerardo Lamas

EXPERIMENTAL HYBRIDIZATION BETWEEN PHYCIODES THAROS
AND P. BaTesit (NYMPHALIDAE). Charles G. Oliver __.__

NOMENCLATURAL CHANGES IN EUCOSMINI (TORTRICIDAE).
Richard L. Brown...

POPULATION STRUCTURE AND GENE FREQUENCY ANALYSIS OF
SIBLING SPECIES OF LETHE. Mark W. Angevine & Peter F.
Brussord.0

THE LARVA OF CRYPTOCALA ACADIENSIS (BETHUNE) (NOCTUIDAE).
Timothy L. McCabe 0

COURTSHIP BEHAVIOR OF THE CHECKERED WHITE, PIERIS PRO-
TODICE (PIERIDAE). Ronald L. Rutowski sae

A NEW TECHNIQUE FOR THE PROSPECTIVE SURVEY OF SEX
CHROMATIN USING THE LARVAE OF LEPIDOPTERA. Wini-
fred Cross & Alison Gill, 0) a

GENERAL NOTES

Temporary range extension and larval foodplant of Dynamine dyonis
(Nymphalidae) in Texas. Joseph F. Doyle III _____-__.-..-------_-__--__-
Oviposition of the butterfly Battus belus varus (Papilionidae). Ailen M.
Young) [220 te
Malfunction of ecdysis allowing imaginal emergence but causing death of
adult hackberry butterfly (Nymphalidae). Raymond W. Neck _.------
Aggregative behavior of Anartia fatima (Nymphalidae) in Guanacaste
province, Costa Rica during the dry season. Allen M. Young ___-_--
Biston betularia, obligate f. insularia indistinguishable from f. carbonaria
(Geometridae). Sir Cyril A. Clarke 2.0.00. a
Brother Apolinar Maria (1867-1949) and his contributions to Colombian
Lepidopterology. Gerardo Lamas ._........-...._.
Weather and the regulation of Hypothyris euclea (Nymphalidae): popu-
lations in northeastern Costa Rica. Allen M. Young ___-----_------_-—

NOTES AND NEWS

Editorial policy statement to contributors 22.0000.)
The James H. Baker collection. J. F. Gates Clarke _............. goa

21
29

37

42

50

Volume 33 1979 Number 2

JOURNAL

of the

_ Leproprerists SOCIETY

Published quarterly by THE LEPIDOPTERISTS’ SOCIETY

Publié par LA SOCIETE DES LEPIDOPTERISTES
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN
Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS

Lv oe 20 July 1979

THE LEPIDOPTERISTS’ SOCIETY

EXECUTIVE COUNCIL

I. F. B. COMMON, President T. SHIROZU, Vice President
C. V. COVELL, JR., lst Vice President JULIAN P. DONAHUE, Secretary
L. A. GOZMANY, Vice President RONALD LEUSCHNER, Treasurer

Members at large:

R. A. ARNOLD J. F. EMMEL C. D. FERRIS
E. D. CASHATT R. R. GATRELLE J. Y. MILLER
R. E. STANFORD ASP. PLATT M. C. NIELSEN

The object of the Lepidopterists’ Society, which was formed in May, 1947 and for-
mally constituted in December, 1950, is “to promote the science of lepidopterology in
all its branches, ....to issue a periodical and other publications on Lepidoptera, to
facilitate the exchange of specimens and ideas by both the professional worker and the
amateur in the field; to secure cooperation in all measures” directed towards these
aims.

Membership in the Society is open to all persons interested in the study of Lepi-
doptera. All members receive the Journal and the News of the Lepidopterists’ Society.
Institutions may subscribe to the Journal but may not become members. Prospective
members should send to the Treasurer full dues for the current year, together with
their full name, address, and special lepidopterological interests. In alternate years a
list of members of the Society is issued, with addresses and special interests. There
are four numbers in each volume of the Journal, scheduled for February, May, August
and November, and six numbers of the News each year.

Active members—annual dues $13.00
Student members—annual dues $10.00
Sustaining members—annual dues $20.00
Life members—single sum $250.00
Institutional subscriptions—annual $18.00

Send remittances, payable to The Lepidopterists’ Society, and address changes to:
Ronald Leuschner, 1900 John St., Manhattan Beach, California 90266 U.S.A.

Back issues of the Journal of the Lepidopterists’ Society, the Commemorative Vol-
ume, and recent issues of the NEWS are available from the Assistant Treasurer. The
Journal is $13 per volume, the Commemorative Volume, $6; and the NEWS, $.25 per
issue.

Order: Mail to Charles V. Covell, Jr., Memoirs Editor, Department of Biology, Uni-
versity of Louisville, Louisville, KY 40208, U.S.A.

The Lepidopterists’ Society is a non-profit, scientific organization. The known office
of publication is 1041 New Hampshire St., Lawrence, Kansas 66044. Second class
postage paid at Lawrence, Kansas, U.S.A. 66044.

Cover illustration: Third instar larva of Limenitis archippus Cramer (Nymphalidae)
preparing to enter winter diapause. The larva is resting on the lip of its hibernaculum
constructed from the basal portion of a chewed tubular willow leaf (Salix babylonica
Linnaeus) covered with silk. In the autumn such larvae begin facultative diapause in
response to decreasing day-length. Original drawing by Mr. George C. Ford, Jr., Graph-
ics Illustrator, Department of Biological Sciences, University of Maryland Baltimore
County, 5401 Wilkens Avenue, Catonsville, Maryland 21228.

Sec

eS eS

JOURNAL OF

Toe LeEepPIpoPTERISTS’ SOCIETY

Volume 33 1979 Number 2

Journal of the Lepidopterists’ Society
33(2), 1979, 77-111

SUBSPECIFIC VARIATION IN BUTTERFLIES: ADAPTATION
AND DISSECTED POLYMORPHISM IN PIERIS
(ARTOGEIA) (PIERIDAE)

S. R. BowpDEN

53, Crouch Hall Lane, Redbourn, Herts. AL3 7EU, England

ABSTRACT. Subspecific variation of butterflies has in the past been attributed
(with doubtful justification) to local ecological factors. In part I of this paper,
main types of variation in wing-color and marking in the Artogeia napi-bryoniae-
melete group are described. Genetic control is relatively simple and known
phenocopies result from environmental abnormalities unrelated to any actual local
conditions.

In part II, selective agents are considered to exercise little or no control over sub-
specific pattern-differences. Physiological adaptation is certain, but visible characters
appear not to be specially adapted to present local conditions.

In part III, the extreme selectionist position is rejected. Phenotypic differences
have arisen historically, often in consequence of the dissection of polymorphisms,
and are maintained by the succession of genetic path-choices in effectively isolated
populations. Territorial changes have involved the retention of pattern-elements in
biotopes far removed, geographically and ecologically, from the places of origin.
Common possession of even superficial characters, when genetically identical, implies
phylogenetic relationship.

Some consequences of range-changes of populations are discussed.

In this paper it will be maintained that wing-pattern (and other
phenotypic) variation among subspecies of butterflies is generally con-
trolled not by present ecology but by factors having an historical aspect.
Arguments will be related particularly to the group Artogeia napi +
bryoniae + melete. The genus Artogeia Verity was formerly included in
Pieris Schrank; however, Kudrna (1974) and Higgins (1975) have cor-
rectly separated it.

For the past hundred years, collectors have amassed long series of

78 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

butterflies and moths showing the variation of species among different
localities, with the expectation that the significance of the diversity would
reveal itself, thus exposing the processes of evolution. I think this expecta-
tion has generally been disappointed. But at the turn of the century it
was believed quite generally that the visible differences between local
races expressed their close adaptation to contemporary environmental
conditions.

This thesis received considerable support from the experiments of
Standfuss (1900) and others on the heat-and-cold treatment of lepi-
dopterous pupae. These appeared to show (inter alia) that central
European races were convertible into the semblance of Mediterranean
races by moderate heat, or into phenocopies of Scandinavian races by
moderate cold. It was supposed that environmental factors themselves
had the power to produce phenotypes better adapted to cope with them,
and that a process rather like that known later as genetic assimilation
(Waddington, 1953) had produced the climatic races. The argument
was in some degree circular, being based on an initial assumption that
the supposed climatic forms were adaptive.

Some local departures from the idealized type were held to be directly
environmental, as when marshy conditions were themselves supposed to
shift the phenotype in a particular direction, such influence not being in-
herited. Other deviations were correctly recognized as selective and
heritable: “chalk” forms of a butterfly or moth were whiter than heath
forms (Poulton, 1890. 157; Ford, 1945: 124; Ford, 1955: 97). There was,
therefore, ample excuse for naturalists who attached what might now
be called ecological import to all local variations.

Their attitude persists among many lepidopterists. Thus Dennis
(1973), in a study of the spotting on the wings of the satyrid butterfly
Coenonympha tullia Miller in a Welsh colony, concluded that “certain
broad spatial features emerge that may prove vital to an understanding of
the ecological pressures on C. tullia,’ when his painstaking analysis had
established that localized demes differ in spot arrangement. I do not
question that local variation occurs, in genotypes as well as in phenotypes;
but it is by no means certain that the groups delimited by marking owe
their visible differences to their present ecologies. Another example is the
strange data concerning spotting on the hindwing of Maniola jurtina L. in
England, claimed as providing ideal material for investigating adaptation
and the action of natural selection (Dowdeswell, 1956). No adaptation
was detected, and the basis of natural selection remained quite un-
certain, “presumably of a physiological nature.”

The significance of wing-pattern is a matter of natural history, re-
quiring in the first place no very specialized techniques for its appraisal.

VoLUME 33, NUMBER 2 79

Indeed, the replacement of the chase by genetic experiment, and even
the application of statistical methods, have not in most cases explained
geographic variation at specific or subspecific levels though we are able
to talk more precisely about it. Nevertheless, I will attempt to reassess
the position—the present position—in relation to one group of pierid
butterflies. This is a very narrow foundation on which to erect any
general principles, but it may be adequate to invalidate the extreme
neo-Darwinian dogma.

Part I. MorpH SYSTEMS IN THE ARTOGEIA NAPI SPECIES-GROUP

In the species-cluster Artogeia napi L., A. (n.) bryoniae Ochsenheimer
and A. melete Ménétriés, we are faced with groups of almost conspecific
taxa, with wing ground-colors varying from nearly pure white to deep
golden yellow and to a tawny ochre, carrying melanic markings varying
from intense and extensive to evanescent. The ground-colors are due to
related pterins, and the melanic pattern has a common basis in all the
populations.

The wing-pattern variation of the napi-bryoniae group in Europe was
meticulously described and figured by Miller & Kautz (1939). Their
treatment of the Asiatic and American taxa was very incomplete and
included few or no illustrations. Indeed, I know of no comprehensive
study of the Nearctic members of this group, nor have I myself all the
material necessary for such an undertaking. Artogeia melete and some
associated Asiatic species were the subject of illustrated papers by Shel-
juzhko (1960-69). The present work is not concerned with the characteri-
zation of taxonomic entities as such, nor will descriptions be in greater
detail than is required by subsequent discussion. The accompanying
figures (Figs. 1-40) are far from illustrating the total range of variation
encountered in the superspecies: nor do they include any forms that
can fairly be regarded as “aberrations.”

It will be convenient to consider background colors (sections 1, 2), the
markings of the hindwing underside (section 3), and upperside markings
(sections 4, 5). Finally the reflection of ultraviolet light, invisible to
vertebrates, will be dealt with in section 6.

These groupings of characters do not exhaust those which might be
considered to have a fairly simple polymorphic basis. For example, in
many Artogeia taxa there is normally a small orange-yellow lunule or
streak near the base of the hindwings, below. Furthermore, in some
Asiatic species, but not others, the forewing discal cell is invaded below
by black scales (character “nigrosparsa”). These appearances will not
be considered further here.

80 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

1. The sulphurea Schoyen polymorphisms!

European A. napi subspecies typically have a white upperside; the
underside of the hindwings and of the forewing apices (i.e. those parts
exposed when the insect is at rest) are yellow in ground-color. The re-
mainder of the forewing underside is white. In the male the yellow is
usually rather paler than in the female.

In the rare variant formerly bred by Head (1939; see also Bowden,
1954) the white pigment is replaced by lemon-yellow, which may be
very brilliant if the adult develops at a high temperature. When this
yellow is paler, it can be seen that the hindwing underside remains
brighter than the upperside: that is, the typical pattern persists. This
form occurs equally in both sexes, and is recessive to wild type.

Another sulphurea allele appears to be responsible for Thompson’s
pale yellow, a form which is fairly distinct in the female, but often almost
indistinguishable from wild type in the male. It also is recessive, but is
dominant to Head’s form. It is uncertain how commonly the gene or
gene-combination determining it occurs in England and in other parts
of Europe. A genetic study was made by Bowden (1961), but further
work on a wider basis is desirable.

Whereas Head’s bright yellow does not occur regularly in any natural
population, a form indistinguishable from Thompson’s, and with the
same dominance relationships, is typical of A. (napi) marginalis Scudder
of Oregon. It may occur also in certain other American populations, but at
present it is not known whether it is anywhere in polymorphic balance
with the white form (Bowden, 1970).

In various bryoniae populations of the eastern Alps (and probably
elsewhere) a dominant gene in some of the individuals prevents the
development of the lemon-yellow (sepiapterin) color but does not affect
the ochreous color which many females show (see section 2). The re-
sulting male phenotype (f. subtalba Schima) is white above and below,
but nearly all the corresponding females have hindwing undersides which
remain ochreous, though their lemon tinge has been lost. The frequency
of this gene (another allele of the sulphurea series: Bowden, 1963) is
locally as high as 0.25 in the alpine region and the system is there truly
polymorphic. |

In those European napi populations in which the great majority of
females are white above, the subtalba morph is virtually unknown. Nor
does it appear in the arctic ssp. adalwinda Fruhstorfer. The extent and

'Infra-subspecific names (such as sulphurea) are not italicized, but are given the author’s name
when first used. The names of characters, if new or not well known, are put in quotes when first
used, but not afterwards.

VoLUME 33, NUMBER 2 81

mode of its known occurrence in mainland Asia (e.g. Amur) is un-
fortunately still obscure.

In the eastern Nearctic species A. virginiensis Edwards a subtalba
gene (also dominant and an allele of sulphurea: Bowden, 1966) is fixed.
Here males as well as females possess a little ochreous pigmentation,
which prevents a conspicuous white underside in either sex. On the other
hand the sympatric P. (napi) oleracea Harris has a light lemon-yellow
underside in both male and female, and we have not yet encountered a
subtalba form in it.

Artogeia melete melete (Japanese) appears to lack sepiapterin in both
sexes, but here again males as well as females have considrable ochreous
pigmentation (especially on the hindwing underside). This could be re-
garded as a subtalba form, but it differs genetically from Schima’s
original subtalba, being apparently recessive when hybridized with
typical napi (Bowden, 1975).

A. japonica, described by Shirézu (1952) as a subspecies of napi, is
specifically distinct from both A. (n.) napi and A. melete (Bowden,
1978a). In appearance it approaches very closely the artificial hybrid
between these two species, but it is of subtalba form and again the
subtalba is recessive to napi wild-type. In this species the underside of
the male is white, not ochreous.

A. dulcinea Butler and some other far eastern taxa possess subtalba
forms, but in all these cases it is still uncertain whether a balanced poly-
morphism exists.

2. The female form flava Kane

As mentioned above, in some (particularly alpine and arctic) popula-
tions the wings of the female (but not of the male) are tawny or ochreous
yellow. The responsible pigment (erythropterin, possibly with other re-
lated compounds) is present not only on the upperside but also on the
hindwing underside, though not normally on the underside forewing
disc. Occasionally even this area is more or less ochreous: in ssp.
neobryoniae Sheljuzhko such specimens are not rare; they occur also
in the arctic ssp. adalwinda.

In ssp. britannica Verity of Ireland and Scotland the sex-limited
ochreous variant has wrongly been regarded as an aberration, and named
flava Kane. In fact it occurs regularly in both populations and its ground-
coloration is apparently identical with that typical of adalwinda females.
As it is unrecognizable in the male and even in the female has variable
expression, its genetics have perhaps never been adequately investigated.

The results published by Lorkovic (1962), however, allow the pos-

82 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

sibility that in bryoniae it is unifactorial and dominant, if incompletely
so, the depth of color developed depending presumably upon modifying
genes. The results (1956 and unpublished ) from my own hybridizations
between bryoniae-like subspecies and various white forms are in ac-
cordance with this finding.

The occurrence of ochreous pigmentation in both sexes of A. virginiensis
and A. melete has been mentioned above. Since in these species the pig-
ment switch is not completely sex-controlled, it must in some way differ
genetically from that in A. napi and bryoniae.

The apparent association of flava color with ultraviolet reflectance will
be discussed in section 6.

3. The underside “green veining”

The species-group is characterized by the borders of black or grey
scales on either side of the veins on the hindwing underside. When the
ground-color is yellow, the visual effect is greenish, though this effect
is lost if the black is too dense or the ground-color is white.

For the purpose of comparisons we shall take as standard the black
bordering as it appears in the spring emergence of English A. napi (Figs.
2, 7). In the non-diapause generations the density of the bordering is
much reduced: an extreme reduction appears in A. napi meridionalis
Heyne (Figs. 13, 17) of Mediterranean countries. Curiously, warming
English napi pupae after completion of diapause leads to slightly in-
creased melanic scaling on the underside (Bowden, 1978b). In any one
population the width of the border does not usually vary very much,
though very broad borders (obscuring most of the ground-color) fre-
quently occur in A. napi adalwinda (Fig. 26) and A. n. hulda Edwards.

A rather broad, vaguely defined vein-bordering characterizes A.
virginiensis (Fig. 29) and helps to separate it from the sympatric but
ecologically distinct A. (n.) oleracea. In the Near East flies A. (n.)
dubiosa Rober (= pseudorapae Verity); the better known non-diapause
generations have vein-marking so nearly obliterated as to justify both
its names, but in the diapause generation form suffusa Verity, which
appears to belong to dubiosa, carries very broad vein-stripes.

A. (n.) oleracea (Figs. 32, 34), like A. melete (Fig. 21), has narrow
and sharply defined vein-marking, which in the spring emergence can be
extremely densely black or chocolate-black. In the non-diapause genera-
tion (Figs. 37, 39) the borders remain narrow, but are only slightly
pigmented and often hardly visible. This “acuta” type of vein-bordering,
found also in the Californian A. (n.) venosa Scudder, is dominant over
both European wild type and the suffused bordering of A. virginiensis

VoLUME 33, NUMBER 2 83

(Bowden, 1972). It retains its general character in hybrids repeatedly
back-crossed to European napi, but the “summer” veining is then more
visible than in the pure oleracea gene-complex. The acuta veining of
A. melete (Figs. 21, 23) also is dominant in hybrids (Bowden, 1975).
All these modifications of the underside veining are retained when the
subspecies concerned are bred in England. It should be said that though
the “seasonal” dimorphism is normally a simple consequence of the
diapause/non-diapause alternative, some intermediate phenotypes (on
both surfaces of the wings) can occur in special circumstances ( Bowden,

1978b).

4. The upperside markings

Again it is convenient to regard the markings of spring napi of southern
England (Figs. 1, 6) as normal, as well as most appropriate for the com-
parison of univoltine populations such as A. n. adalwinda and A. (n.)
bryoniae.

In the male (Fig. 1) the forewing discal spot varies in size within the
same subspecies and may be absent, though in this case one or two spots
on the underside of the forewing (Fig. 2) normally remain. The female
marking (Fig. 6) includes on the forewing two discal spots, with often a
supra-discal spot more or less covered by the extension of the apical.
The pre-apical costal mark prominent in A. krueperi Staudinger is some-
times faintly represented, and may be distinguishable from the supra-
discal spot proper (when both are present) by a different degree of
melanization. These additional spots are sometimes apparent in males
also (especially in non-diapause emergences ), but a dark streak along the
hind margin is normally characteristic of the female sex. The dark basal
suffusion is more extensive in the female, whose forewing veins are
lined-in, especially towards the outer margin, where this black scaling
commonly expands to form dark triangles.

On the fore-margin of the hindwing of both sexes there is usually a
black spot, more or less in line with the forewing discal spots but homol-
ogous rather with the apical. Marking along the hindwing veins is vari-
able, but generally light.

In bryoniae-like forms (Figs. 4, 9, 27) the female radial vein-marking
is intensified and broadened, and an additional streak (“Saumstrich” or
“bryo-streak” ) appears outside the middle of the lower (i.e. posterior)
discal spot; basal and marginal dark suffusions may also become ex-
tensive. The combinations of these elements in various proportions are
infinitely varied, so that no two females are entirely alike. Stipan et al.
(1960) attempted the thankless task of completing Kautz’s descriptive

84 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

naming (Miller & Kautz, 1939) of the main possible forms. Development
varies between and sometimes within populations from a mere bryo-
streak to the extereme f. concolor Rober, approached by Fig. 9. The not
uncommon f. “meta” is illustrated in Fig. 5. Clearly the bryoniae pheno-
types cannot be due to a single dominant gene, as has sometimes been
supposed. Apart from modifiers, there may be several linked major
genes involved. Probably only the “radiata” element has been ade-
quately studied genetically, and this is indeed a simple dominant
(Lorkovic, 1962).

In non-diapause individuals (Figs. 3, 8) the spots become more promi-
nent, the radial markings less so. Extreme forms, in this direction, are
found in the summer broods of A. n. meridionalis. Females representa-
tive of this subspecies (Fig. 16) lack even the marginal black streak con-
tinuing the upper edge of the lower discal spot; this streak is always
absent in A. rapae L. (Fig. 20).

Even in summer males, the discal spot can be completely absent (as it
is in Pieris brassicae L.), but this is uncommon in Europe. The aspect
of both sexes is changeable by a varied development of the supra-discal
spot and its confusion with the apical (Fig. 12).

The upperside markings of A. melete (Figs. 22, 24) differ from those
of A. napi by their greater intensity and breadth, which frequently leads
to their confluence. In the male melete the second discal spot appears
regularly.

5. Restricted upperside markings

When over-wintering pupae which have completed diapause are held
at varying temperatures between O°C and the temperature of normal
development (ca. 6-8°C) some individuals are diverted to produce a
“superspring’ form (Figs. 14, 19). Though the forewing apical spot
and the corresponding spot on the hindwing may persist, the discal spot
markings disappear completely or almost so, even from the female and
even on the underside, and the radial viens are more or less blackened
throughout their length. In extreme examples the sexes become alike
(Bowden, 1978b ). This form, which can be obtained in several (perhaps
all) European subspecies, including adalwinda and bryoniae (Fig. 18),
may be called “restricta’—the name referring to a character and not to
any taxonomic entity. Genetically, the treated insects showing it are
quite normal (usually wild type) for their subspecies.

Some Nearctic napi subspecies present a similar or more extreme ap-
pearance even in their natural temperature regime. P. (n.) oleracea may
be quoted: in both diapause and non-diapause generations (Figs. 31-34,

VoLUME 33, NUMBER 2 85

36-39) the napi spot-markings are faint or completely absent, the apical
marking is weak and there are no spots even on the underside of the
forewings; on the upperside even radial marking is slight or absent.
In oleracea-napi hybrids this genetically determined restricta behaves
as a simple recessive, linked with the dominant acuta (Bowden, 1972).

Ssp. marginalis (Figs. 35, 40) is slightly less restricta in phenotype
than oleracea; females usually show the black streak on the forewing
hind-margin, the adjacent discal spot (rather weakly) and sometimes
a very weak anterior discal spot. Crosses with oleracea suggest, how-
ever, that the same restricta gene is present in both subspecies.

Ssp. venosa Scudder of California lacks the restricta character, though
acuta is present; in this it resembles some taxa of eastern Asia.

6. Ultraviolet patterns

In recent years many workers have studied the reflection of ultra-
violet light by butterflies. Among the Pieridae, the yellow butterflies
such as Gonepteryx ( Nekrutenko, 1964), Colias ( Ferris, 1973) and Eurema
(Ghiradella et al., 1972) show ultraviolet-reflecting patches in the male
only, and these are structural, a complex lamellar system on the ridges of
the scales producing optical interference.

In Artogeia, on the contrary, the males usually remain dark (as do
the females also in A. napi napi), but in many subspecies the female
upperside reflects more or less strongly. This is so particularly in those
European taxa in which the female ground-color is ochreous, but A. (n.)
macdunnoughii Remington of Colorado and A. melete have nearly white
females which are rather brightly reflective. Experiment shows that
absorption by varying concentrations of leucopterin in the wing-scales
and (in the females) structural reflection are responsible for the dif-
ferences (Bowden, 1977).

Part I]. ADAPTATION AND SELECTIVE NEUTRALITY

From the time of Darwin (1859) on, the basic idea connecting variation
with evolution has been that of adaptation—often a vaguely formulated
idea, leading to tautological arguments whose circular character easily
escaped detection.

In some balanced systems, such as the melanisms (Kettlewell, 1973)
and the mimetic complexes (Sheppard, 1961), there are indeed good
reasons for supposing that primary selective advantages are involved and
often the basis of selection can be proposed with confidence. The selec-
tion is intraspecific, acting on the individual, so that in particular circum-
stances one allele is favored up to an equilibrium level. Population

S6 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

10

EXPLANATION OF FIGURES

Fie) Dio N. Lie cps: 6) 65 2X2 6 N L. ups. 2 ’46 Cornwall
2 N- Ti. wns, 6. "6922 7 N L. uns. 9 69w%25
oN) $2 ups: 3 ’50iHents: 8 N_ S. ups. 9 ’44 Essex
4A DB ie ups... 3 73b 10 9 B. Li ups. 9 765rag
Do (BN)? S. aps, «2/5210 10 BU. uns. 2 “7eb25

Abbreviations: A = ssp. adalwinda (Lappland), B = ssp. bryoniae (Swiss), (BN )°
=F. hybrid 9B x 6N, D=ssp. meridionalis (Corsica), J=sp. & ssp. melete
(Japan), K=ssp. neobryoniae (Karnten), KA =F; hybrid 9K x ¢A, M=ssp.
marginalis (Oregon), N = ssp. septentrionalis (England), O = ssp. oleracea (New
Hampshire, U.S.A.), Ra=sp. & ssp. rapae (England), V=sp. & ssp. virginiensis
(Connecticut, U.S.A.). L.= from diapause pupa, S. = from non-diapause pupa,
ups. = upperside, uns. = underside. Figures and italic letters following sex-signs are
individual identifications.

VOLUME 33, NUMBER 2 87

i 16

12
i7
13 18
14 19
15 20
Bieta Soups: ¢ '64d°*45 GD eSa ups. 2 640525
Peewee swups: 6 ‘64d°'5 i Doe uns. (OMe OAd 74
ieee suns: ¢ '64d**11 Lo) By wisps. O77 2AI4
14 NE... ups: ¢ ’708 [97 IN| Ee-ups: 2 270¢'6
ease. ups. 2 53924 20) Ravisssups. © 164 Herts:

Editor's Note: Figures reduced to 0.88 of original size.

density is not usually determined by such selection: it can seldom
happen that the population-level of an insect is fixed by the bird predation
which is selecting the cryptic or mimetic adult.

There has been a widespread opinion, following Fisher’s (1930) argu-
ment, that genes having a neutral effect, that is on balance neither ad-
vantageous nor disadvantageous, are necessarily extremely rare. All

88 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fic. 21 J L. uns, 6 71s 95 A LL. ups. 6 sake
7g Jott 96 A I. uns. 6 veddsa

aes esa cs id 97 A L. ups. 2 '34d95
23s Onseuc ea 2yin 0 98 VV Te) ups) 6 ameue

29 Vo i. uns: oe

2A Sees Sey ye ai 30 V 'L. ups) S7Gbgio

Abbreviations: A = ssp. adalwinda (Lappland), B = ssp. bryoniae (Swiss), (BN )*
=F. hybrid 9B x 6N, D=ssp. meridionalis (Corsica), J=sp. & ssp. melete
(Japan), K=ssp. neobryoniae (Karnten), KA =F: hybrid 9K xX 6A, M=ssp.
marginalis (Oregon), N = ssp. septentrionalis (England), O = ssp. oleracea (New
Hampshire, U.S.A.), Ra= sp. & ssp. rapae (England), V = sp. & ssp. virginiensis
(Connecticut, U.S.A.). L. = from diapause pupa, S. = from non-diapause pupa,
ups. = upperside, uns. = underside. Figures and italic letters following sex-signs are
individual identifications.

VoLUME 33, NUMBER 2 89

3|
32
33
34
35

mews Ol. ups. ¢ ’650°16 S677 Ow Se ups 6 “CoO22
SOs uns, 3) ’650°13 Si On Seaunss aoa O3O27
somO We ups. 2 ’650°12 38 O S. ups. 2 ’65030
548 Ome uns. 2 ’650'39 Se ©) S, tik, © GHORY
Sone ve ups. 2’ ’66M"***10 40 M S. ups. 2 ’680***9

Editor’s Note: Figures reduced to 0.88 of original size.

present characters of subspecies must then be expected to have been
positively selected, that is, to be “adaptive.” When visible characters
appear to have no conceivable selective value, appeals to the pleiotropy
of genes are commonly made to escape the contradiction: a correlated
physiological modification, perhaps affecting behavior or environmental
response, is assumed. When the phenotype in question varies locally in
frequency it is necessary to think of a covert character, controlled by the

90 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

gene concerned, which would have different optimization in the various
localities. Such characters are not entirely nonexistent, but how are
they to be identified? And as Robson & Richards (1936: 274) remarked,
“It is no use to smuggle these facts of specific differentiation into the
proof of natural selection by an appeal to ignorance, or by an assumption
of correlation.” In fact an hypothesis supported by a supposition of
pleiotropy has no standing until at least the close genetic linkage of a
known associated effect has been independently established—calculation
of a statistical correlation is seldom convincing by itself.

Moreover, as modifying genes were supposed ultimately to render
unfavorable characters recessive (Fisher, 1929), so they should also
reduce the penetrance or expressivity of such characters in pleiotropic
systems. The appeal to pleiotropy itself admits that the positive selection
of a gene need not imply positive selection for all the characters con-
trolled by the gene; visible characters may then well be neutral. And if
some of the characters controlled by a pleiotropic gene can be non-
adaptive, these seems no reason why other phenotypic characters should
not be so. Thus the argument from pleiotropy may be set aside as for
our purpose irrelevant. Our concern here is with visible characters: have
they been positively selected by external forces, or are they maintained
as externally neutral factors in some well-balanced genetic complex?
One should allow that even a character originally fixed by selection may
thereafter become effectively neutral in a variable environment. For
plants, Stebbins (1974) has pointed out that many complex adaptive
structures have been retained long after the strong selective pressures
that were required to establish them have ceased to exist (e.g. in self-
fertilizing Leguminosae and apomictic Compositae). We shall suggest
that whether or not genes can rarely be neutral, characters controlled by
them quite commonly are. This can hardly be denied (Ford, 1945a: 78).

Curiously, no one seems to maintain that characters which have become
specific ones are fixed by environmental agencies. Clossiana selene
Schiffermiller and C. euphrosyne L. are now sympatric and the differ-
ences in their patterns owe nothing to present local conditions. How
are the wing-markings of Nymphalis antiopa L. determined? Normalizing
selection is working on ancestral genotypes, probably with no visible
response to ecological conditions. If we consider two Artogeia species
which have become sympatric, say A. napi and A. rapae, it is clear that
at some unknown time these had a common stock, which has since been
subject to repeated splitting. It would be difficult to believe that present
ecology determines their pattern-differences; why then should we be-
lieve that it does so for allopatric ssp. napi and ssp. oleracea?

In the last few years Fishers “dogma,” which led to various para-

VoLUME 33, NUMBER 2 91

doxical results in the hands of other mathematicians, has indeed ap-
peared less cogent. “Mutations” rather suddenly came to include not
only conspicuous alterations of the phenotype but also changes in pro-
teins, detectable by electrophoresis, and even replacement of one genetic
codon by a synonymous one. Changes in proteins in the course of evolu-
tion were investigated by their analysis in nearly and distantly related
taxa (Goodman et al., 1971). King & Jukes (1969) concluded, rightly
or wrongly, that at the molecular level most evolutionary change might
be due to selectively neutral mutations and genetic drift. Kimura & Ohta
(1971) also maintained that the protein polymorphisms known from
electrophoretic data were transient phases in the random-walk fixation
of neutral mutations. Theoretical difficulties were exploited, or ex-
plained away, by the mathematical population-geneticists (e.g. Karlin
& Nevo, 1976) though, as M. Nei said in discussion, “We cannot make a
general inference about nature from a study of a specific mathematical
model.” But evidently a belief that neutral mutations may have a part
to play, at the allozyme or even at the morphological level, no longer
involves the scientific ostracism of its holder.

ADAPTATION IN ARTOGEIA

In what sense are the characters which visibly differentiate related
populations of Artogeia adaptive? What common agencies could be
expected to exert selective pressure, on the basis of visible characters?

Firstly, the physical environment itself might be directly operative, or
might interact with physiological processes.

Secondly, predation by birds, lizards and various arthropods might
be influenced by the procryptic or apostatic patterns of individuals.

Thirdly, such predation might be reduced, in unpalatable groups, by
_ warning coloration.

Fourthly, intraspecific pairing might be facilitated, and interspecific
pairing discouraged, by the butterflies’ recognition only of the appropri-
ate patterns.

These, the more obvious possibilities, will now be considered.

1. Physical environment

It must first be remarked that the genus Artogeia, as well as many
Satyridae, has been particularly influenced by the climatic changes of
the last few hundred thousand years, and it may well be that some
distribution-changes are too recent for close approach to genetic equi-
librium. As will be seen, this may affect the validity of some arguments.

992 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Subspecies in this group differ in mean size, but so do the two or three
generations annually within a subspecies. Particularly common is a slight
dwarfing in later summer emergences, attributable to the effect of
drought on food-plants. In general, however, napi populations from the
Mediterranean region tend to produce large adults. Levins (1968) has
remarked that contra-gradient variation is common among invertebrates—
those in hotter, drier regions may be “genetically” larger to attain the
same actual size in their own environment. One effect of the conflicting
factors has been to produce some extreme individuals in these climates
(cf. Holl, 1914; Warren, 1970). Inter-subspecific dimensional compari-
sons are therefore best made between insects reared together in captivity;
otherwise it is easy to draw incorrect conclusions.

The physical environment certainly exercises control over other
physiological adjustments, and also modifies the expression of subspecific
wing-characters in each individual; but convincing evidence for its selec-
tive influence on these last has been almost entirely lacking.

The absorption of solar radiation and its conversion into heat assists
poikilothermic animals to become active. Watt (1968) has studied the
thermoregulation of six species and subspecies of Colias and shown ex-
perimentally that the greater melanization of the underside in the higher
altitude populations helps them to approach their optimum temperature
by appropriate orientation behavior.

Two circumstances may be thought to favor the suggestion of a similar
role for the melanic marking of A. napi, as has indeed been postulated
by Shapiro (1977) in the case of A. n. venosa. Firstly, the underside
veining of the summer broods is always lighter than that of the spring
emergence of the same subspecies. Secondly, the summer broods of the
Mediterranean populations are less extensively melanized below than
those of central and northern Europe. However, the interrelation between
the directly environmental (diapause/non-diapause) and the inherited
factors in these differences makes it difficult to estimate the likelihood
of their adaptive origin.

Nor is climatic basis of the melanic cline very certainly established.
Descimon & Renon (1975), working with the satyrid butterfly Mela-
nargia galathea L. in France, found a melanization gradient exactly con-
trary to that presented by A. napi populations; nevertheless they did not
fail to find an eco-climatic rationale.

Again, the explanation usually given for the ochreous color and melani-
zation of ssp. bryoniae and adalwinda females is that the pigmentation
serves to increase absorption of radiation and so enables these females
to reach a relatively high body-temperature at high altitudes and lati-
tudes. However, the males too need to attain a certain temperature for

VOLUME 33, NUMBER 2 93

successful fertilization; also Pontia callidice Hiibner at even higher allti-
tudes lacks ochre pigment and is not particularly heavily marked.

In the Colias polymorphism involving the form alba (or helice Hiibner )
the frequency of the paler pterin pigmentation actually increases towards
the far north (Hovanitz, 1944), although Watt (1974) showed that in
C. eurytheme Boisduval orange females heat up more rapidly in sun-
light than alba females, as would be expected from their greater ab-
sorption of blue light. But Lorkovi¢c & Herman (1963) confirmed the
unequal viabilities of the C. croceus Fourcroy genotypes at different
temperatures in experimental conditions which presumably eliminated
any color effect. Descimon’s (1976) conclusion would make the color-
difference an incidental consequence of the diversion of nitrogenous
compounds from pigment formation to egg-nutrition in the helice form.

Petersen (1952, 1955) has shown a correlation of ochreous pigmenta-
tion with relatively heavy melanic marking in Artogeia populations of
mixed phenotype (ssp. neobryoniae). Since both characters are more
or less completely dominant, this correlation might be, at least in part,
a consequence of hybridization of a genotype producing both with a
genotype producing neither. There is no evidence of linkage (Lorkovic,
1962; Bowden, 1956 and unpublished ): the expected recombinations take
place in the experimental F, and the corresponding phenotypes are to be
found also in the wild. In the Fennoscandian territories occupied by ssp.
adalwinda, bicolorata Petersen and napi, Petersen (1949) found that the
clines for female ochreous ground-color and for heavy melanic marking
did not altogether coincide.

Lorkovic (1962) concluded that “we can say very little about possible
pleiotropy of the color genes” and that if there were no correlation with
physiological and ecological characteristics there must be a directly
genetic selective advantage. Local variation of the frequency of morphs
_was earlier considered to be a reliable indication of the action of selection.
Perhaps this begged the question a little.

How the subtalba balance is maintained, ecologically or genetically
(by heterozygous advantage), is not yet known. Bowden (1967) dis-
proved the suggestion of lethality of the homozygote, but could not
exclude, in the varying genotypes of the eastern Alps, the possible associa-
tion of subtalba with linked genes which reduced the number of homo-
zygous females in some broods.

Ford (1964: 291) says, “The bryoniae complex of genes and characters
evidently fits the butterfly for life at high altitudes and latitudes... .”
but “It seems unlikely that the same genes are responsible for bryoniae
throughout the whole of its discontinuous distribution.” Since the dif-
ferent elements of the bryoniae phenotype (see above, under “Upper-

94 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

side markings”) are unequally developed in different localities less than
500 km apart, this is certainly true to some extent, but his further discus-
sion would suggest that bryoniae is not a monophyletic group, similar
characters having instead been selected, by similar environmental con-
ditions, in local stocks. This is difficult to believe: Petersen (1958) was
so convinced of the common origin of the female patterns in ssp. bryoniae
and ssp. adalwinda that he was prepared to allot the latter subspecies to
the “species” bryoniae, although it belonged reproductively to napi.

Robinson (1971) offered the following on f. subtalba: “In either napi
or bryoniae the subtalba form occurs as part of a polymorphism, but in
Pieris virginiensis the form is actually the wild type. In the Nearctic
habitat of virginiensis the conditions so favour subtalba that the wild
type found in either napi or bryoniae is completely lacking.” Such an
ecological “explanation” explains nothing; besides, it is partly contradicted
by the napi-wild-type coloration of the sympatric A. (n.) oleracea.

A further objection to facile ecological derivations is that in this group
of butterflies each population has some freedom to select its own habitat
in a locally varying environment. The phenotype, in fact, comes to choose
the kind of selection to which it will be exposed (Waddington, 1967). It
is even reasonable to suppose that if the American and European napi
populations were wholly interchanged they would survive by this means,
each with its own markings, though the oleracea colonies in Europe
would not necessarily persist in the precise localities now occupied by
A. napi.

2. Cryptic patterns

It seems obvious enough that the veined pattern of the underside con-
tributes much to crypsis at rest, as do the mottled patterns of some other
pierids (Pontia, Euchloe). The usually yellow ground-color, whether
lemon or ochre in tint, adds to the success of this effect. It is notable
that in most of those morphs or subspecies which lack the lemon-yellow
sepiapterin, the females or even both sexes have available another pig-
ment (essentially erythropterin) which produces a pale orange (ochre-
ous) effect (Bowden, 1966). The basic pattern of the underside can
thus well be called adaptive.

The differences among the subspecies probably cannot: all variants
evidently work sufficiently well. If one compares oleracea with napi, it
is difficult to suppose that the former benefits from its more conspicuous
veining in spring, since this veining practically disappears in non-diapause
emergences. Can its American niche demand veining more definite than
the European in spring, less definite in summer? Have these butterflies,
in all localities, less need for crypsis in summer than in spring? It is

VOLUME 33, NUMBER 2 95

difficult to answer these questions in the affirmative. The differences
are surely not cryptically adaptive.

The subtalba polymorphism (where it exists as such) could be main-
tained by selection of apostatic forms (Clarke, 1969), in this case by pre-
ferential predation of the locally commoner form of male. In the female,
with its greater reliance upon crypsis, the morphs are not very unlike
(relative to the variation within each morph) and even the most dis-
criminating predators would hardly distinguish between them. But this
possible mechanism may in any case be precluded by aposematism.

3. Warning coloration

Poulton (1890: 185) remarked, “The colours which produce . . . the
greatest effect, upon the eye of an insect-eating vertebrate, are black and
white... .”

It has been stated that Pieris and Artogeia (especially perhaps P.
brassicae ) are distasteful to birds, and that the white color is aposematic.
I have indeed observed that sparrows (Passer domesticus L.) are not
always effectively deterred from taking P. brassicae, and a captive Euro-
pean gecko captured and ate the same insect. Collenette (1935) quotes
numerous anecdotal reports of attack by birds on P. brassicae and A.
rapae (and on an occasional A. napi); in very few was the insect re-
jected after being secured. However, there were some observations of
definite avoidance. Probably attacks are far fewer than a palatable
insect would suffer. _

Indeed, Rothschild et al. have recently established (Marsh & Roth-
schild, 1974) that all three British whites feeding on crucifers contain
toxic substances which would render them relatively noxious to birds,
though only P. brassicae is sufficiently so to have evolved aposematic
rather than cryptic coloring in the larval stage. The mimicry is Mullerian
rather than Batesian, since not only is the supposed model, brassicae,
usually less numerous than rapae and napi, but these mimics are them-
selves toxic.

Certain further facts do support the synaposematic hypothesis:

(i) The close pattern-resemblance of Artogeia rapae to Pieris brassicae,
an insect otherwise so different as to merit its full generic separation
(cf. Warren, 1961, Kudrna, 1974).

(ii) The full development of the same pattern in A. napi only in the
Palaearctic region, where A. rapae flies naturally.

(iii) The formation of a similar Mullerian (?) group in eastern Asia, by
A. melete, A. japonica and A. napi, again with A. rapae. Here the

96 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

superficial resemblance of japonica to melete is very close indeed
(A. melete is toxic—Rothschild, Marsh & Bowden, unpublished).

However, one would say that, to predators, differences in the black
upperside patterns exhibited by A. napi subspecies in most different
localities would probably be irrelevant, neither adding to nor detracting
from any deterrent effect. And napi does vary independently of its
“models” within the European and east Asiatic regions. A. ergane Geyer,
which belongs to the wider napi grouping, is a much closer mimic of
A. rapae.

If, as seems likely, the white butterflies evolved originally from yellow
or orange ones not unlike Colias, the aposematic effect of white may
have contributed to its success in replacing orange. But orange-and-
black, too, is a warning coloration, and Collenette (loc. cit.) quotes a
case of house sparrows taking white butterflies but neglecting Colias.

The pale lemon yellows found in rare variants in Europe would be
aposematically inferior to both orange and white, but if so the fixation
of “pale yellow” in the ssp. marginalis of Oregon requires a special ex-
planation which is lacking at present.

White is conspicuous in nearly all Pieris and Artogeia, except that in
the bryoniae-like subspecies the females are dark and show little white.
Since the females’ individual protection is the more important to the
species, there is an implication here that the males’ gain from their white-
ness is by no means indispensable.

4. Pair-formation

The experiments of Eltringham (1933), well summarized by Ford
(1945b), showed that male butterflies recognize their females at a dis-
tance by their color. One can observe that male whites will approach
any white butterflies, including other males, though at close quarters
scent cues and behavior patterns become more important.

On the other hand Petersen and collaborators found that searching
male bryoniae are attracted (from a distance) to white napi females,
even in marked preference to their own dark females (Petersen, 1963).

Petersen's experiment needs repeating and extending, but his reported
result could be expected from the apparently dull female coloration, if
bryoniae react equally to ochreous and white stimuli. But there is a
complicating factor: these tawny females reflect ultraviolet light,
whereas most white and lemon-yellow males and females do not. As has
long been known (Lubbock, 1899) insects not only distinguish colors
but also see the near ultraviolet so that these “dark” females might ap-
pear brightly ultraviolet. Such reflection is not in fact confined to

VOLUME 33, NUMBER 2 97

ochreous individuals but is just as strong in certain “white” females, for
example those of A. (n.) macdunnoughii.

The absorption of ultraviolet light by the males’ wings renders them
“colored” to the females and may help to release pairing behavior,
though in conditions of proximity scent is almost certainly predominant.

Obara (1970) claims to show that ultraviolet reflection is the only
signal exciting sexual behavior in the A. rapae male, and that markings,
size and shape of the wings (and even scent) are all irrelevant. This
conclusion cannot, of course, be extended to A. napi, many of whose
females do not reflect ultraviolet.

Adaptation for sexual selectivity between species is given by the ex-
istence of any specific difference readily perceptible by the insects con-
cerned; neither form is “superior” to the other, advantage for both being
obtained merely by magnifying the difference.

What, then, should be our assessment of the influence of local selective
pressures on the visible characters which distinguish related Artogeia
populations? Wide variability of essentially the same designs would
itself suggest either that the variants are approximately neutral in effect,
or else that a number of separate optima exist, each related to the local
environment. The latter alternative does not accord with observed simi-
larity of phenotype over geographic areas comprising a range of biotopes.

For example, the extensive biotopes of napi and oleracea in Europe and
America vary greatly, as from moorland to deciduous forest, and the
heterogeneity is relatively small-scale, with Nearctic and Palaearctic
habitats overlapping in character. Clearly the sharply veined phenotype
is not being selected by positive environmental pressures peculiar to the
New World.

Many subspecies of A. napi can be regarded as exerges in the sense
of Verity (1925, 1953). In his quoted case of Mellicta athalia athalia
Rottemberg and M. a. celadussa Fruhstorfer, the chief observable dis-
tinction is in the male genitalia, but fully fertile interbreeding neverthe-
less occurs along the subspecific frontier, introgression producing a
hybrid zone varying between 50 and 160 km in traceable width ( Higgins
& Riley, 1970; Higgins, 1975; Guillaumin & Descimon, 1976). Should one
suppose that one form of genitalia is adaptive in the conditions of north-
ern and central Europe, the other in Spain and Italy?

Similarly, within the A. napi group androconial scale shapes vary; we
can see no advantage in this, though as Lorkovic (1970) has pointed
out, variation (if it occurred) in the male scent disseminated might con-
tribute to sexual discrimination; at the specific level the lemon-verbena
scent of A. napi is quite distinct from the faint sweetbriar of A. rapae

98 JouURNAL OF THE LEPIDOPTERISTS SOCIETY

(Ford, 1945b). The seasonal androconial differences described by
Warren (1961-1967) also must surely be nonadaptive.

We have shown above (see also Bowden, 1978b) that both upper-
and underside markings in the A. napi group are subject to direct en-
vironmental modification; differences between post-diapause and non-
diapause adults often exceed those between distinct species. Nor is there
sufficient reason to suppose that these phenotypic departures are pri-
marily adapted to special seasonal requirements, though advantage may
be taken of them when they favor physiological adjustments. The
restricta marking of oleracea is nearly phenocopied by European sub-
species subjected to an unnatural temperature regime. It is not impos-
sible that the facies characteristic of the various populations are de-
termined by rate-genes which produce differential realization of the
elements of the basic pattern, and the seasonal differences are brought
about quasi-automatically. Current utility of a character need not imply
adaptive origin; the well documented cases of pre-adaptation (cf. Huxley,
1942) are sufficient evidence of this.

There is no need to doubt that these Artogeia are well attuned to their
rather wide niches, and that this adaptation was achieved through natural
selection. Local adaptation, however, seems to be physiological—in
respect of such matters as voltinism, temperature-tolerance, etc. This
will prevail, whether or not every element of pattern is optimal for the
necessary processes.

In the subspecies of A. napi, it appears that ecological conditions
may now exert little beyond normalizing selection on the genes controlling
some visible characters. Elements of the wing-pattern, if they are thus
unresponsive to the particular milieu, can be given the greater weight in
taxonomy at the species-level and below. The general principle is ex-
emplified even by genitalia and androconia (cf. Robson & Richards,
1936: 299; Ford, 1945a: 79).

Limits of Selection Theory

As Lewontin (1974) remarks, we cannot know the overall importance
of balancing selection by demonstrating that it exists—of course it
exists. The problem is, what proportion of observed genic variation is
maintained by selection? The school of ecological genetics would at-
tempt to solve the problem by establishing and quantifying the selective
forces involved in as many cases as possible. But it is doubtful whether
anyone has succeeded in measuring the net fitnesses of genotypes for any
locus in any species in any natural environment (Lewontin, 1974: 236).
Indeed, even attempts to assess reliably the mean relative advantages of

VOLUME 33, NUMBER 2 99

alleles in experimental conditions are defeated by the difficulty of secur-
ing a representative set of individuals for the trial (e.g. Bowden, 1967).
And necessarily, where selection is frequency-dependent, the establish-
ment of equilibrium proportions implies that the genotypes present are
equally fit at that equilibrium. It is to equilibria that published mathe-
matical treatments generally refer—and they are almost powerless to
involve real time. Fitness may depend not only on the character of the
remainder of the genome, but also on the gene’s own frequency in the
population (Dobzhansky, 1970). Hartl & Cook (1976) claimed to show
that purely random selection could maintain genetic polymorphisms even
when the expected fitness of each genotype was the same.

Part III. DissEcrep POLYMORPHISM IN SUBSPECIATION
Dissection of Polymorphisms

Many distinctive characters, as we have indicated, are controlled by
single genes, and if the differing patterns coexisted in one population—as
they sometimes do—would constitute a classical genetic polymorphism
(Ford, 1940, 1961).

Lewontin (1974) has said, on the basis of observations on Drosophila,
that “the overwhelming preponderance of genetic differences between
closely related species is latent in the polymorphisms existing within
species. This must be even truer of those between subspecies, but we
frequently find that within Artogeia populations there is in fact little
heterozygosity in respect of the distinctive characters that we recognize—
hence indeed their taxonomic value.

I shall not attempt to discuss here the general question of the main-
tenance of heterozygosity in Artogeia populations. In very few insect
species are sufficient data available—certainly not yet in A. napi. Studies
of enzyme polymorphisms, how they are balanced and how dissected
within and between its populations, may soon provide such data and
at the same time suggest phylogenetic relationships. Gene-frequencies
close to fixation are very insensitive to selection and have hardly any
implications about the recent past; on the other hand a gene-frequency
close to 0.5 offers chiefly information about the recent past and es-
sentially none about the remote past (Lewontin in Moorhead & Kaplan,
1967).

Consider a former polymorphic species extending its range and in the
course of ages being divided by geographic catastrophe or otherwise into
isolated subspecies. In subspecies b one morph may become fixed, in
ssp. c the other, while ssp. a retains the original polymorphism. Even
if a later becomes extinct, the dissected polymorphism (Bowden, 1970)

100 JoURNAL OF THE LEPIDOPTERISTS SOCIETY

is still traceable in b and c. This must have been a common sequence
for the differentiation of characters of slight as well as of marked adaptive
significance. Ford (1945a: 83) quotes the striking case of the arctic fox,
Alopex lagopus L.

In the sulphurea polymorphisms of A. napi, the alleles (four if one
includes the near-extinct bright yellow) probably became established
by three successive dimorphisms of this kind, though one cannot yet
determine their order.

There is of course a general if not universal condition for the main-
tenance of polymorphism (while it lasts), that the population over a
period should be at least as viable with two morphs as it would be with
either alone. When the morphs have become fixed in allopatric popu-
lations or even distinct species, it seems that at some time and place
this condition has ceased to be satisfied.

But this overlooks some consequences of the variation within the
precursor population. Since its genotype is not uniform, the samples
taken from it to produce b and c will have differed, and if these founder
populations have been small, either originally or in subsequent fluctua-
tions, some genes included in a may not be represented at all in b (or c).
Moreover, as May (1976) reminds us, replicate lines from the same
initial population, kept in identical conditions, can reach very different
limit-compositions, as a result of statistical accidents early in the breeding
program.

The so-called fixation of one morph theoretically does not involve the
permanent elimination of the other, even in a single population. Re-
current mutation may be expected to cause its persistence as the hetero-
zygote at a very low fluctuating level, even if with intermissions. It
might seem that this could permit the reconstitution of the polymorphism
if selection-pressures came to favor it, or even the evolution of a new
genotype around the temporarily rare allele. But the chance of this must
steadily decline and finally vanish.

The irreversibility of evolutionary changes was postulated by Meyrick
(1854), though apparently only for changes of generic or higher rank.
Muller (1939) already concluded, “The determination of the exact muta-
tional path of evolution involves a large element of accident and . . . this
path can never really be retraced, nor paralleled, in a second evolutionary
sequence, nor can the same complex genic system be twice arrived at.”
At any level, the principle follows from the mutual dependence of genes
in their operation. As a mutant at one locus proceeds towards fixation,
other parts of the genotype undergo changes, so that a simple reverse
mutation no longer restores the original condition. For true restoration
a series of reverse changes must take place in the correct order—with

VOLUME 33, NUMBER 2 101

probability approaching zero. An evolutionary event is reversible: an
evolutionary history is not (Dobzhansky, 1970). Thus I believe that
lost characters are rarely restored, and perhaps never with quite the
original genetic mechanism.

While our precursor species remains intact, we have a number of
individuals, occupying the same niche, in competition. But if b and c
are separated geographically, interbreeding ceases and so does repro-
ductive competition. The two daughter gene-pools evolve along separate
historical successions, genotypes in Db now competing only with other
b genotypes; whether c has evolved “superior” genotypes is irrelevant.
Thus it is not certain that b and c are both better adapted in their own
locations: it is possible that they would do equally well if interchanged.

Species appear to have been free to make a wide range of replies to
the same ecological demands. It is remarkable how different organisms
in the same habitat, and even in practically the same niche, utilize such
diverse means of “adaptation” to it (thus of course modifying the niche).
The strength of their defense against the selective tyranny will be ap-
preciated more readily by the botanist (Willis, 1940; Gavaudan, 1967;
Stebbins, 1974) than by the zoologist, who is tempted to endow his
mobile subjects with niches of any complexity that his theories demand.
“Within any gene-pool there exist several . . . alternative gene combina-
tions that might adapt the organism to any new environment. The
particular adaptive combination that will become established will depend
largely upon the nature of the gene-pool already present” (Stebbins,
1974). If a population could be divided equally between two separate
identical biotopes, it is probable that even then two identical subspecies
would not be formed.

Only in its marginal habitats is the adaptation of a species critical for
its survival. Butterflies are able to accomodate variation in the repro-
ductive success of the “same” population by factors of ten or twenty to
one (much larger fluctuations have been quoted for other organisms);
beside such ratios selective advantages of even five or ten percent re-
lated to a particular imaginal character (even fertility) are quite in-
significant if they apply between separated populations. This does
not mean that, within each, selection does not still act on the character—
merely that the apparent alternatives are not in competition, if the taxa
concerned are no longer interbreeding. Allopatric differences cannot
be maintained by selection between them.

The relative stability of a subspecific genotype results from its achieve-
ment of an adaptive peak, almost any small departure from which will
involve disadvantage. In a widespread species there are many such peaks,
and although some are “higher” than others, a particular population will

102 JoURNAL OF THE LEPIDOPTERISTS SOCIETY

normally be unable to pass from one to another. Only where a cline
exists peaks are not separate but form, as it were, an elevated ridge,
allowing free variation in the clinal character.

In any one environment, at any time, there may be several adaptive
peaks accessible to a population. Once it is on a slope, selection will
drive it upwards, but it cannot climb two separate peaks at once unless
it is no longer panmictic. The initial difference which determines the
peak to be climbed is at that stage probably nearly neutral.

Nor does the peak genotype’s integration imply that any one character
has optimum individual adjustment. This is clear enough where distinct
“island” subspecies occur in the same climatic region; examples are
well known in Erebia (Satyridae). In that genus the different wing-
patterns may tell us something about past relationships, but very little
about present ecology.

“Les corps inanimés ne dependent pas du temps. Les corps vivants lui
sont indissolublement liés. Chez eux, aucune structure ne peut étre
détachée de lhistoire” (Jacob, 1970). Whatever is, is good—but it may
not be the best.

Origins, Range-changes and Hybridization

In recent years systematists (e.g. Brundin, 1972; Darlington, 1970;
Croizat et al., 1974) have been debating newly formulated ideas on
biogeography. The accepted view, deriving originally from Darwin, had
been that a species arose at a particular place and if successful spread
therefrom. Adams (1902) provided criteria for the determination of
centers of origin, the most important being the “location of the greatest
differentiation of a type.” Matthew (1915) deduced that the most ad-
vanced species would be found at the center of origin and the most primi-
tive or conservative in peripheral areas. However, Hennig’s (1966)
“phylogenetic systematics,” sometimes identified with cladism, implies
that speciation is always by division of a pre-existing species-stock (e.g.
by geographic catastrophe ) to produce vicariant populations which then
evolve separately in their own areas: these are recognizable as belonging
to one monophyletic group by their common possession of derived
(“apomorphous’) characters. One of Hennig’s rules states that species
with the most primitive characters are found in the area earliest occupied
by the group—thus contradicting the formerly accepted view.

Some of the contestants approached bigotry in their philosophical dis-
cussions, and the subject can now be advanced only by consideration of
particular examples.

Too little is yet known about Artogeia. Using Adams’ criterion, the

VoLUME 33, NUMBER 2 103

genus was given an origin in central Asia, and writers such as Miller &
Kautz (1939) supposed that species and subspecies spread therefrom
to the north, west and east, reaching North America via the Bering
area. These taxa, then, differed even at the start of their journeys and
retained their characters long after arrival. On the other hand, Hennig-
Brundin principles would suggest that, while any secondary characters
held in common are evidence of common descent, the characters dis-
tinguishing the subspecies arose only in the various localities after con-
tacts were interrupted, and represented the results of local adaptation.

Both alternatives are too simple. The evolution of Artogeia, in Pliocene
and. Pleistocene times, has been complicated by relatively rapid altera-
tions of climate and by fluctuating sea levels affecting the extent of ex-
posed ice-free land. Time-scales are such that, in many if not all cases,
territories must have changed since the differentiation of the taxa—a
process almost indistinguishable in its results from conventional dispersal.
One has to suppose that major continental drift in the Atlantic area
occurred too early to affect Artogeia, but some seas, and some mountain
ranges, are younger.

Even in a region as small and well-known as the British Isles, there is
still some uncertainty about the relationships of the populations of A.
napi. Its position is simpler than that of Aricia agestis Denis & Schiffer-
muller/artaxerxes Fabricius (Hgegh-Guldberg & Jarvis, 1969), in that
it is unnecessary to propose any specific separations, but there are
parallelisms. Verity (1911, 1916) placed the dark-female butterflies of
Ireland and Scotland in his ssp. britannica, while naming those of England
as ssp. septentrionalis. Warren (1968) took the Scottish populations
(but not the Irish) as belonging to the “species” adalwinda, and named
them ssp. thomsoni; his view was based on a strange argument concern-
ing their androconial scales. In 1970 Lees recorded the existence of a
univoltine race in Yorkshire at about 300 m elevation. Thomson (1970)
found that androconial thomsoni extended as far south as Yorkshire, but
was everywhere at least partly bivoltine.

An adequate discussion would be out of place here. There seems little
doubt that the Irish and Scottish populations, which regularly include
ochreous females and darkly marked ones, derive partly from adalwinda-
like insects which occupied the Channel area during the last stages of the
Wirm (Wisconsin) glaciation. In the short post-glacial period A. napi
napi has invaded Britain from the South and its introgression has
progressed so far that in southern England the modern ssp. septentrionalis
hardly differs from the nominotypical subspecies. Even in Scotland
there are probably now no pure relict “adalwinda.” Indifferently as to
whether they carry subarctic genes or not, all members of the Scottish

104 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

demes are adjusted physiologically to their present biotopes. Where and
how the original separation of napi adalwinda and n. napi took place is
quite unknown.

In North America, mountains made conditions complex during the
retreat of the ice, and true relict populations may exist—ssp. marginalis
and perhaps macdunnoughii?

Even in the last ten thousand years or so insects whose predecessors
were separated at much more remote periods have come into renewed
contact while still interfertile. The resulting zone of secondary inter-
gradation (Mayr, 1942) is commonly marked by a proliferation of forms
in unstable polymorphisms. The instability is long-period and so cannot
be established with certainty. Probable examples are to be found in the
Artogeia “subspecies” flavescens Wagner and neobryoniae; the zones of
introgression are quite deep, suggesting that the gene-complexes of the
original constituent races were basically similar. Indeed without such
similarity the gene-expression might be disturbed (Kettlewell, 1965,
1973). Variation in chromosome number within a population, as in A.
(n.) neobryoniae (Lorkovi¢, 1970) may or may not lend support to the
hypothesis of hybrid origin, but the significance of B-chromosomes in
the A. napi group is still uncertain.

Warren (1966-69) has derived many apparently normal Artogeia
species or subspecies from processes of hybridization (necessarily in
secondary contacts). As he writes, we have for example oleracea and
other Nearctic subspecies taking their origin from A. narina Verity x
A. dulcinea Butler (Warren, 1968). These supposed parent species must
be understood as ur-narina and ur-dulcinea, because they may since have
undergone as much change as the “hybrids” evidently have. If we reject
Warren's unsatisfactory androconial criterion of hybridism (Bowden,
1971) there is no obvious reason to exclude the possibility of an ur-
oleracea contemporary with the precursors of narina and_ dulcinea.
Hennig (1966) seems to take the view that when a new species has split
off from an existing stem-species, the stem-species also must be taken as
changed. I am not sure that this will always be so, when new species
are formed from peripheral subspecies. Nor is the possibility of reticulate
evolution (i.e. close hybridization) to be altogether excluded: there
may sometimes have been more than contact-line introgression. We
should try to infer phylogenies for our species from the evidence that
we can collect, but a large element of conjecture will have to be tolerated.

The Alaskan A. (n.) passosi Warren may well be the oleracea x
hulda hybrid that Warren (1968) supposes, though not reproductively
isolated from either oleracea or hulda. In the secondary contacts of the
napi populations of Scandinavia, the British Isles and southern Europe

VoLUME 33, NUMBER 2 105

the distribution of characters in the conjoined populations has perhaps
become clinal, with no definite limits to the mutual introgressions. If the
characters are adaptive or have become linked to adaptive ones, local
optimization may control the proportions of the morphs in a frequency-
cline, but otherwise recognizable distinguishing characters reflect origins
rather than adjustments to the varying environment. Stepped clines
may be better considered as arrays of micro-subspecies, unless appropri-
ate partial barriers can be found located at the steps. See, however,
Endler (1977).

Ford (1949) treats British Coenonympha tullia populations as a pres-
ently interrupted cline, and postulates some flow of genes between many
of them even today, but in fact his Merioneth xX N. Scotland crosses
showed disturbances almost too great for a mere cline of 700 km. Even
a continuous cline, with its assumed equilibrium conditions, may not
always be distinguishable from an historical pattern originating from
range-change by differing routes.

Clarke (1970), commenting on the Cornish boundary phenomenon in
Maniola jurtina (Creed et al., 1959), suggested that two races of the
species might have been involved, though Ford (1975) rejected this view
with extreme scorn. Nevertheless, an “area effect” (Cain & Currey,
1963), of adaptation to inherited genotype rather than to external ecology,
may indeed be operative.

CONCLUSIONS
Special

1. There is no evidence that the difference in Artogeia napi between
narrow American underside veining and broader European veining is
adaptive. Both crypsis and optimum radiation-absorption can be secured
with either arrangement of black scales.

2. In the “summer” emergence of various subspecies there may be
adaptive adjustment of the degree of underside blackening for the most
favorable balance between cryptic and thermoregulatory functions.

3. The subtalba polymorphism, controlling production of sepiapterin
or failure to produce it, is dissected in the Artogeia napi species-group.
In certain populations, particularly in the alpine regions of the Old World,
the polymorphism is a balanced one, at least on a medium time-scale.
Though proportions of the morphs vary locally, no ecological basis is yet
known and explanations may be historical.

4. Form subtalba may have reached the bryoniae butterflies from an
ur-melete or related Asiatic taxon, but if so changes have since occurred
in its genetic control.

106 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

5. Other alleles at the sulphurea locus will repay further study in a
wider range of populations in the three continents concerned. The pale
yellow forms are stabilized in at least one (probably relict) subspecies.

6. The significance of the bryoniae female marking remains uncertain.
The taxa carrying it were in the past conspecific with other branches of
the napi stem. Miller & Kautz (1939) considered it beyond question
that these taxa represented the conservative (even archaic) descendants
of the Pliocene, probably subtropical, insect. This is no more than con-
jecture, but not unreasonable; if it is correct, what we have to account
for are not the dark females but the white males of bryoniae and both
sexes of napi.

7. Artogeia adults are cryptic at rest, probably aposematic in flight.
Local differences in pattern may be irrelevant in both respects.

General

8. Overt differences between allopatric populations of the same species
or species-group need not be supposed in every case to have any present
adaptive significance. Some varying characters shown in the course of
a life cycle may be well adapted to the habitat, but many are approxi-
mately neutral. Competition between their holders can only be internal
to a population. The central weakness of the extreme selectionist position
may lie in its assumption that a determinable fitness is associated with
any particular gene.

9. The permanence of subspecific characters (and _ still more of
specific) depends on their tendency, once fixed in the genetic sense,
to become irreversible in practice.

10. Characters that are now nonadaptive may have been evolved at
a remote period in a locality far removed from the present habitat.
Biogeography is not merely a branch of ecology. Ecological fallacy
results from treating populations as stationary objects of local selective
pressure; instead, they should be allowed a more active role, with every
evolutionary quasi-random “choice” determined in some degree by the
influence upon the genome of previous choices. The saying “Evolution
always occurs somewhere else” is nearly true.

11. When the “same” character arises independently in two stocks
by parallel evolution, parallelism will probably be incomplete and a
genetic difference can sometimes be found, to alert the investigator.

12. Otherwise, the common possession of the same “nonadaptive”
characters permits tentative conclusions on phylogenetic relationships.
The barrier to the reversal of a genetic change is an increasing one,

VOLUME 33, NUMBER 2 107

allowing the established form to be taken as indicator (though not as
final proof) of the affinity of subspecies which possess it.

13. Discontinuities of pattern-distribution can often be used to de-
limit demes whose past histories have not coincided.

14. Genetic histories do not repeat themselves, whatever the ecological
pressures.

Murray (1972), discussing the work of Kimura & Crow and others on
electrophoretic alleles in Drosophila, etc., says: “There are loci which
are monomorphic in all populations, there are loci with rare variants, and
there are loci which so variable that no ‘wild type’ can be identified.
However, there is a kind of differentiation that is conspicuously absent,
i.e. a pattern of variation with the fixation of different alleles in different
localities. This last pattern is the expected outcome of allelic neutrality.”

Publications on the allozymes of Artogeia are awaited. But is this
missing pattern not to be found in the visible characters of this group?

In one sense, all important characters of the napi group are adaptive,
and are maintained by soft selection (Wallace, 1968). But the visible
differences between the various taxa, with which this paper has been
concerned, are not primarily adaptations to the present environments
but derive from historical “accidents” affecting neutral or nearly neutral
polymorphisms in the distant past. As far as can be seen, there has
always been more than one way forward for a subspecies. Even under
strong selective pressures there are alternatives, and each option taken
modifies the choices which will be presented in the future.

ACKNOWLEDGMENTS

I am indebted to an anonymous referee and to Dr. Bryan Clarke for
valuable criticism of earlier versions of this paper.

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33(2), 1979, 112-123

NOTES ON THE LIFE CYCLE AND NATURAL HISTORY OF
BUTTERFLIES OF EL SALVADOR. III C. HISTORIS ODIUS
AND COEA ACHERONTA (NYMPHALIDAE-COLOBURINAE )

ALBERT MuyYSHONDT, JR. AND ALBERTO MuySHONDT

101 Avenida Norte #322, San Salvador, El Salvador

ABSTRACT. A complete photo-illustrated report on the early stages of Historis
odius (Fabricius) and a partial of Coea acheronta (Fabricius) are presented, which
reveals similarities between the two species suggesting a very close relationship.
Both species utilize the same foodplant in El] Salvador, Cecropia mexicana
(Moraceae), whose close relative, C. peltata, has been reported as foodplant in Brazil
by some authors under the vernacular name “Embauba.” The placement of these
two species in the Coloburini is questioned and some striking larval similarities with
Smyrna blomfildia and S. karwinskii (both also questionably placed in the Coloburini)
and with Pycina zelis (placed among the Vanesiini) are pointed out. Larvae of
Historis odius are subject to heavy parasitization mostly by tachinid flies and one case
of nematode parasitization, (Mermis sp.), is recorded. Historis odius is by far more
abundant in El Salvador than Coea acheronta and covers a wider range of altitudes.

This article presents information on the life cycles and natural histories
of the two largest species of butterflies included in the Coloburini:
Historis odius (Fabricius) and Coea acheronta (Fabricius).

The first time we saw the eggs of Historis odius was 6 October 1970
when our good friend Viktor Hellebuyck captured and papered a female.
Some 15 eggs were deposited by the female while still alive inside the
envelope. The only information we had at the time about the foodplant
of the larvae of H. odius was a name found in Seitz (1920) and later re-
peated by Ehrlich & Ehrlich (1961): “Embauba.” Knowing the hosts
of related species, we placed some of the eggs of H. odius on the food-
plant of Colobura dirce (L.), Cecropia mexicana Hms. (Moraceae), and
some on the foodplant of Smyrna blomfildia Fabricius and S. karwinskii
Hiibner, Urera caracasana (Jacquin) Grisebach (Urticaceae). In due
time the eggs hatched and the larvae placed on Cecropia ate it for three
days but then died, while the ones on Urera all died without feeding.
Since then we have been able to rear the species several times from eggs
and larvae collected in the field, on Cecropia mexicana.

Coea acheronta has proven to be more difficult. In 18 years of collect-_
ing in the country, only one adult has been captured and only one larva
has been collected. The larva was taken in the fifth instar on Cecropia
mexicana also and reared to adult.

i . asl ate 4 2 . ‘ ‘ ‘ I uy 7
Specimens of eggs, larvae and pupee of H. odius were preserved in alcohol as were the larval

skin and pupal shell of Coea acheronta and ;
Sarasota, Florida, USA, ie iemomanen:

VoLUME 33, NUMBER 2 113

LirE CYCLES
Historis odius

Egg (Fig. 1). Spherical, light brown with 23 lighter vertical ribs, armed with thin
spinulets or thick, short setae running from base of egg to % of the side where
they merge into a circular arrangement of two rows of hexagons, leaving a large
octagon around micropyle. Diameter ca. 1.5 mm. Hatches in 5-6 days.

First instar larva (Fig. 2). Head dark brown with sparse setae. Body cylindrical,
gray before feeding, olivaceous afterwards, with transverse rows of thin, short, light
setae. Ca. 3.5 mm long. Moults in 3-4 days.

Second instar larva (Fig. 3). Head dark brown, surrounded by a row of short,
lateral spinulets and short, thick epicranial horns armed with rosette of short spines
distally. Body dark brown with whitish transverse markings on thoracic segments,
white spots subspiracularly from Ist to 5th abdominal segments and white transverse
stripes caudad. Short white scoli with light spines, furcated distally, on all segments
except Ist thoracic. Ca. 8-9 mm long. Moults in 3-4 days.

Third instar larva (Fig. 4). Head squarish with indented epicranial suture, low
triangular frons with no visible adfrontal areas. One spine standing at each side
of superior end of epicranial suture; a short, thick, rough-surfaced horn on each
epicranium. Each with terminal rosette of short spines: two directed forwards, two
backwards and one vertical; lateral row of spines diminishing in size down to ocelli;
another spine frontally, another slightly lower and a final one at center of the arch
of the ocelli. Color is variable: either all dark brown or dark brown with orange spot
under epicranial horns and another around ocelli. In the latter case, spines on head
capsule and tips of horns are also orange. Body ground color dark brown, masked
thoracically and from 6th abdominal segment caudad by transverse whitish stripes.
Scoli and spines whitish also. Spine arrangement: Ist thoracic (T-1) with dark
cervical shield having subdorsal scolus with two terminal spines and dark spiraculum
having one simple spine above and slightly behind; T-2 and T-3 with 5-furcate
subdorsal scolus, simple supraspiracular spine very close to anterior border of seg-
ments, one bifurcate supraspiracular scolus and one simple subspiracular spine. First
abdominal segment (A-1) with one dorsal 4-furcate scolus, one 4-furcate supra-
spiracular scolus; 2-furcate scolus and a simple spine anterior to spiraculum. A-3 to
A-7 as A-2 but lacking subdorsal scoli. A-8 with two dorsal scoli, two supraspiracular
scoli, one simple spine and one 2-furcate subspiracular scolus. A-9 with one supra-
spiracular 5-furcate scolus directed backwards. A-10 with 2-spined anal fork. All
scoli and spines light yellow except orange dorsally from A-2 to A-6. Grows to 17
mm in 5 days.

Fourth instar larva (Fig. 5). Head as in third instar but larger. Body ground
color brown with transverse whitish stripes on all segments, ventral surface reddish.
Grows to 38 mm in 2-4 days.

Fifth instar larva (Fig. 6). Head about 8 mm wide, same colors as in third
instar. Body with white or light yellow stripes on red ground color. Scoli reddish on
white-striped morph, yellowish on yellow-striped morph. Ventral surface reddish or
orange. Spiracula and anal fork, black. Grows to 70-72 mm in 6-7 days.

Prepupa. Slight general discoloration and reduction in length. Duration one day.

Pupa (Figs. 7-9). Lateral aspect: very flat, abdominal 10, 9 and 8 thickening
abruptly to A-7 then gradually to A-3 (thickest part of body), with indentation on
A-1 to T-3, then humping dorsally on T-2 then reducing to head, which terminates
with two round, dorsally incurved, partially apposed cristulae about 10 mm long.
A-1 with dark wart at meson, another subdorsally and one supraspiracularly. From
A-2 to A-7 one dorsal 4-furcate spine, one supraspiracular dark wart, and from A-3
to A-7 another spiracular wart. Cremaster located ventrally on a peculiar formation
on the last abdominal segments. Ventral aspect: abdominal segments thicken bluntly
from cremaster, about the same width up to wingcases, which are slightly thicker

114 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

up to about head level; angles abruptly narrowing to the root of head cristulae. Color
pinkish brown with black-tipped, reddish spines and red rings around cristulae.
Dimensions: 50 mm long, 13 mm dorsoventrally, 12 mm laterally at widest points.
Adult emerges in 10-14 days, females being slower than males.

Adults (Figs. 10-13). No noticeable sexual dimorphism. Forewing with convex
costal margin, with projected angled apex, S-shaped outer margin to round tornus,
straight inner margin. Hindwing with slightly convex costal margin, rounded
outer margin to somewhat acute anal angle; inner margin with strong fold. Dorsal
color on forewing, orange basally with black border from mid-costal margin to apex,
descending to tornus, then back to mid-inner margin forming a rounded dark area.
White subapical spot close to costal margin. On hindwing the basal orange is strongly
reduced, the rest being dark brown with a light brown thin border along outer margin.
Ventral color of both wings, mostly light brown with black lines parallel to body.
No “eyes” present. Subapical white spot also visible on underside. Body concolorous
to respective dorsal and ventral colors of wings. Eyes and antennae reddish-orange.
Wing span varying from 75 to 90 mm, females usually larger than males. Total
developmental time, 36—45 days.

Coea acheronta

Fifth instar larva (Figs. 14-15). In all respects like Historis odius except for the
color. Head basically red with black horns and an irregular spattering of black spots
of various sizes. Body mostly dull black with three saddle-like yellow markings ex-
tending to subdorsal area on abdominal segments 2, 4 and 6. A broken brown band
along spiracula sprinkled with tiny white dots. Spines arranged as in H. odius, of
light color on scoli concolorous to body zones. Spiracula black with light brown
borders. Ventral surface white. Single specimen grew to 65 mm.

Prepupa. No color change. Lasted for one day.

Pupa (Figs. 16-18). Very much like H. odius, the only differences are the size
and relative position of head cristulae, which are shorter, not incurved or apposed,
but slightly divergent. Total size about the same as H. odius. Adult emerged in 9
days.

Adult (Figs. 19-20). Same color pattern dorsally as H. odius with five additional
white spots on forewing from mid-costal margin to mid-outer margin, and a black
spot on hindwing near anal angle. Differences in shape are noticeable; apex not
projected, but rounded, smoother outer margin, small tail continuing Mz vein of hind-
wing. Antennae orange with black tip. Wingspan 80 mm.

NATURAL HISTORIES

Most of our observations on the early stages and adults refer to Historis
odius although what we have learned about Coea acheronta corresponds
very Closely. The adults are large, robust, pugnacious and fast flying.
Males exhibit very strong territorial defense behavior. They perch on
tree trunks, usually on hilltops, and chase any intruding flying insect as
well as birds. At times several conspecific males favor the same area,
leading to “ritual fights” between two or more males. We call them
“ritual fights” because the opponents seldom, if ever, actually come in
contact in their rapid circumvolutions even when there are many males
involved. This kind of fighting we have noticed also in many other species
even not related to Historis: Phoebis spp. and Hamadryas spp., with the

VoLUME 33, NUMBER 2 115

Figs. 1-6. Historis odius: 1, egg, ca. 1.5 mm diameter; 2, first instar larva, ca.
3 mm long, resting on perch; 3, second instar larva, ca. 8 mm long, on bared vein; 4,
third instar larva, ca. 15 mm long; 5, fourth instar larva, ca. 38 mm long; 6, fifth instar
larva, ca. 72 mm long.

additional “clicking,” and many Lycaenidae. These fights end when one
or more of the rivals tires and returns to their perching trees. At times
there are males of different species perching in the same area. Usually
there is no interaction unless one of them flies over another perching
male. In such cases, the intruder is chased and flies back to his perch
without any resistance.

H. odius, males in particular, are attracted to mud puddles, fermenting
fruits (either on the ground or even while still on the tree) and to sap
oozing from tree wounds. We have never seen them on any kind of ex-
crements. At times they come to rest on humid roadsides or ravines.

116 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Figs. 7-13. Historis odius: 7, pupa, dorsal aspect, 50 mm long; 8, pupa, lateral
aspect; 9, pupa, ventral aspect; 10-11, female, dorsal and ventral aspects, 90 mm
wingspan; 12-13, male dorsal and ventral aspects, scale in cm.

Females ready to oviposit, visit mid-sized Cecropia trees, alighting on
mature leaves, usually on the underside, and repeatedly act as if deposit-
ing an egg without actually doing it. It is after several such performances
that they drop from the underside of a leaf to the upper surface of a

VoLUME 33, NUMBER 2 IIL 7

siti

I

ee
Genesee

FP PE

Figs. 14-18. Coea acheronta: 14, fifth instar larva, 65 mm long; 15, close-up of

larval head; 16, pupa, dorsal aspect, 48 mm long; 17, pupa, lateral aspect; 18, pupa,
ventral aspect.

lower one. There the female deposits a single egg, always in the central
area of the leaf. Several eggs might be thus deposited on the same tree,
but never on the same leaf.

The newly hatched larva eats the upper part of the shell, down to the
lateral ribs, leaving the rest untouched, and then crawls to a border of
the leaf. It starts feeding at the end of a vein, usually at the tip of a lobe,
and bares the vein while constructing a resting perch with frass pellets
agglutinated with silk. Unlike other species that make resting perches,
H. odius leaves the perch naked, without hanging pieces of leaf tissue

118 JoURNAL OF THE LEPIDOPTERISTS SOCIETY

from it. During the first instar the larva may drop from the perch when
disturbed, but is able to return to it by means of a silk thread that it
produces while falling, as is done by many larvae of Heterocera. First,
second and third instar larvae stay all day on the perch, moving from it
only to feed. In this respect they behave differently from all Limen-
itidinae, which stay on the perch while feeding so that the bared vein
by the perch is extremely long and has a barrier of dry bits of leaf tissue
all around the eaten portion. H. odius may make more than one perch
during the first three instars, but uses one after the other.

The perch is abandoned starting with the fourth instar and the larva rests

VoLUME 33, NuMBER 2 119

conspicuously on the upper surface of the leaf, which suggests that the
species may be protected against predation by the foodplant components.
The larva may also be protected by its sharp spines even though these
do not have stinging properties. The larvae of H. odius are subject to
parasitization, mostly by tachinid flies. We found one larva that was
killed in the fourth instar by a parasitic nematode, Mermis sp. The
nematode was many times longer than the larva (ca. 170 mm versus 31
mm ).

When ready to pupate the larva of H. odius wanders about the plant
until a leaf or stem is chosen; it weaves a silk pad. The larva attaches
thereon its anal prolegs and hangs for a day with head and thorax
ventrally incurved after voiding its digestive tract. The large, con-
spicuous pupa hangs freely from its cremaster and reacts vigorously
to the least provocation with lateral wigglings. It has no protective
coloration or shape as does Colobura dirce (Muyshondt & Muyshondt,
1976).

The adult emerges rapidly from the pupal shell and hangs from it
while expanding its wings and ejecting a rust-colored meconium. Some
45 min later the butterfly is ready to take flight. The energetic and ex-
tremely fast rustling flight of H. odius is very similar to Prepona and
Archaeoprepona spp.

The foodplant, Cecropia mexicana, is discussed in our paper on Colo-
bura dirce (Muyshondt & Muyshondt, loc. cit.). Historis odius is found
from sea level to about 2000 m elevation, the same as Colobura dirce.
Coea acheronta occurs from 800-1200 m.

DISCUSSION

The generic name Historis was established by Hubner in 1819 with
Papilio odius Fabricius as the type-species. Coea also was created by
Hubner in 1819 with Papilio acheronta Fabricius as the type-species.
According to Hemming (1967) both genera are valid. Yet there are
several authors (e.g. Smart, 1975; Riley, 1975; Barcant, 1970; Brown &
Heineman, 1972; Ehrlich & Ehrlich, 1961) who place odius and acheronta
in Historis. Still others, old and modern (e.g. Lewis, 1974; Seitz, 1907-
1924; Schatz, 1885; Reuter, 1896; Doubleday et al., 1849) place the two
species under separate genera with the complication that some use the
generic name Aganisthos Boisduval & LeConte for Historis and Megistanis
Doubleday for Coea! According to Hemming (1967) Aganisthos was
estblished in 1834 with the type-species being Papilio orion; however,
orion and odius are now considered conspecific. From the same source
we see that Megistanis was established by Doubleday in 1844. Its type-

120 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

species is Papilio cadmus Cramer, which is the same as acheronta, yet the
name Megistanis is also valid.

The similarities we have found in the early stages of odius Fabricius
and acheronta Fabricius suggest that both species are congeneric, re-
gardless of the evident differences between the adults. However, we
leave the resolution of such matters to the taxonomists.

Doubleday et al. (1849) compared Historis (as Aganisthos) with
Siderone and Prepona, yet he considered Coea (as Megistanis) to be the
“American representative of the typical Nymphales.” Boisduval (1870)
stated: “Ce genre est bien voisin des Prepona, dont il ne differe guere
que par la forme des ailes. Les chenilles, d’apres le dessin que nous avons
vu, sont tout-a-fait semblables.” (This genus is closely related to Prepona,
the only difference being the wing-shape. The caterpillars, as per a
drawing we have seen, are completely alike.) This statement is far from
reality as can be easily seen by comparing the illustrations in this article
with those in our papers on Prepona omphale octavia (Muyshondt,
1973) and Archaeoprepona demophon centralis (Muyshondt, 1976).

The early stages of H. odius have been at least partially described as
early as 1886 (Miiller) from some drawings made by Burmeister (1878)
of alarva and pupa. Also accordingg to Miiller, Dewitz reported the food-
plant to be Cecropia peltata L. Stoll (1791) presented the drawing of a
Historis larva thinking it was an Anaea. In modern publications there are
some sketchy descriptions (Hayward, 1931; Howe, 1975; Riley, 1975)
and one more elaborate (Brown & Heineman, 1972) that in some respects
matches our own. Hayward (loc. cit.) mentions “Embauba” (as a genus! )
as the foodplant of Historis. Of Coea acheronta there is only an assump-
tion made in Brown & Heineman (loc. cit.) of a larval and pupal de-
scription made by Wolcott based on information received from E. G.
Smyth. Riley (1975) mentioned the same thing. Ours seems to be the
first complete, illustrated description of Historis odius and the only ac-
count (admittedly incomplete) of Coea acheronta.

Comparing the early stages of these two species with the early stages
of the other local species currently included in the Coloburini, we see
that there are hardly any points in common with Colobura dirce (Muy-
shondt & Muyshondt, 1976) except for the use of the same foodplant.
Compared with Smyrna blomfildia and S. karwinskii (Muyshondt &
Muyshondt, 1978) they show superficial resemblance during the larval
stage, but not as eggs or pupae. Both Smyrna use Urticaceae, not
Moraceae, as foodplants. The early stages of Pycina zelis Godman &
Salvin resemble those of Historis odius and Coea acheronta more than
Smyrna. The egg is about the same size and has the same shape but a
different number of ribs (32 in Pycina versus 23 in Historis); the larva

VoLUME 33, NUMBER 2 pall

also has the same general aspect but different coloration. Although the
pupa of Pycina, according to drawings supplied by Dr. A. H. B. Rydon
of pupal shells at the British Museum (Natural History), is not like
Historis, Pycina adults do have the same color pattern dorsally as does
Coea acheronta, the underside being more like Smyrna. Pycina zelis feeds
on Urticaceae as does Smyrna.

The eggs of Historis odius and Pycina zelis (probably of Coea also)
have spinulets or setae that are more or less thin. The only other eggs
we have seen with setae, even if much thinner, are the eggs of various spe-
cies of Adelpha, Biblis and Mestra. The shape of these eggs though have
nothing in common with the eggs of Historis or Pycina. Adelpha eggs,
like all Limenitidinae, are “pineapple” shaped and covered all over with
hexagonal cavities. Eggs of Biblis and Mestra are slightly cone-shaped
with vertical ribs reaching close to the micropyle and covered with a
profusion of thin setae.

In Historis odius we again find the similarity noticed in many other
species feeding on plants reputed to be poisonous or at least containing
noxious or strongly aromatic compounds; the larvae wander openly on
the leaves of the foodplant and have gaudy colorations that suggest un-
palatability or some other type of protection against predation. The
adults also have showy color combinations, at least dorsally, many in-
cluding orange, a color that has been associated with predator-deterrent
properties in insects. Yet these larvae are heavily decimated by parasites,
especially Diptera and Hymenoptera. Interested readers may refer to
Muyshondt (1974 & 1975) for our hypothesis regarding the preference
of parasitic Diptera and Hymenoptera for hosts protected from predation
by foodplant derivatives. Regarding the parasitic nematode, Mermis sp.,
the only other time we personally found it affecting lepidopterous larvae
was in 1964 when an early outbreak of Trichoplusia ni (Hubner) oc-
curred on cotton in the central area of El Salvador. Many larvae were
found dead, still clinging to the leaves, with the nematode abandoning the
host through the anus. Then in 1972 our good friend Steve Steinhauser
gave us a larva of Quinta cannae Herrick Schaffer which produced three
nematodes. The nematode was many times longer than the host in every
case.

ACKNOWLEDGMENTS

We feel greatly indebted to Dr. G. L. Godfrey for reading, criticizing
and suggesting improvements to our article. One of us, A. Muyshondt, Jr.,
thanks the Zoologische Staatsamlung in Miinchen and the British Museum
(Natural History) for allowing him to photocopy many old works that

122 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

helped us document our presentation. We are grateful to Dr. A. H. B.
Rydon for supplying us with several valuable publications for the same
purpose.

LITERATURE CITED

Barcant, M. 1970. Butterflies of Trinidad and Tobago. Collins. London.

BorspuvaL, J. A. 1870. Considérations sur des Lépidoptéres envoyés du Guatemala
a M. de lOrza. Oberthur et fils. Rennes.

Brown, F. M. & B. Herneman. 1972. Jamaica and its butterflies. E. W. Classey
Ltd., London. xv + 478 p.

BurMEIsTER, H. 1878. Description physique de la République Argentine. Vol. 5
(Lepidoptera). Buenos Aires.

Dovus.epay, E., J. O. Wesrwoop & W. C. Hewrrson. 1849. The genera of diurnal
Lepidoptera. Vol. 2. London.

Eureuicu, P. R. & A. H. Enruicu. 1961. How to know the butterflies. W. C.
Brown Co., Dubuque, Iowa. 262 p.

Haywarp, K. 1931. Los Nymphalidos Argentinos. Rev. Soc. Ent. Argentina 4,
nos. 1-3.

Hemminc, F. 1967. The generic names of the butterflies and their type-species.
(Lepidoptera: Rhopalocera). Bull. British Mus. (Nat. Hist.), Entomology
Suppl. 9. 509 p.

Howe, W. H. (Ed.). 1975. The butterflies of North America. Doubleday & Co.,
Inc., Garden City, N.Y. xiii + 633 p.

Lewis, H. L. 1974. Butterflies of the World. G. Harrap & Co., London. xvi +
103 p.

MULLER, W. 1886. Stidamerikanische Nymphalidenraupen. Versuch eines natiir-
lichen Systems der Nymphaliden. Zool. Jahrb. Zeitschr. Syst., Geogr., Biol.
der Thiere 1; 417-678.

Muysuonpt, A. 1973. Notes on the life cycle and natural history of butterflies of
El] Salvador. I. Prepona omphale octavia (Nymphalidae). J. Lepid. Soc. 27:
210-219.

1974. Notes on the life cycle and natural history of butterflies of El

Salvador. V.A. Pyrrhogyra hypensor (Nymphalidae-Catonephelinae). J. New

York Entomol. Soc. 82: 162-172.

1975. Notes on the life cycle and natural history of butterflies of El

Salvador. VI A. Diaethria astala Guérin. (Nymphalidae-Callicorinae). J. New

York Entomol. Soc. 83: 10-18.

1976. Notes on the life cycle and natural history of butterflies of El
Salvador. VII. Archaeoprepona demophon centralis (Nymphalidae). J. Lepid.
soc. 30; 23-32.

Muysuonpr, A., Jr. & A. MuysHonpr. 1976. Notes on the life cycle and natural
history of butterflies of El Salvador. IC. Colobura dirce L. ( Nymphalidae-
Coloburinae). J. New York Entomol. Soc. 84; 23-33.

1978. Notes on the life cycle and natural history of butterflies of El
Salvador. IIC. Smyrna blomfildia and S. karwinskii (Nymphalidae). J. Lepid.
soc. 32: 160-174.

Reuter, E. 1896. Uber die Palpen der Rhopalocera. Helsingfors.

Riney, N. D. 1975. A field guide to the butterflies of the West Indies. Quad-
rangle/The New York Times Book Co., New York. 224 p.

VoLUME 33, NUMBER 2 123

ScuHatz, E. 1885. Die Familien und Gattungen der Tagfalter. Lowensohn in
Furth (Bayern).

Serrz, A. (Ed.). 1907-1924. Die Grossschmetterlinge der Erde. Vol. 5, Die
Amerikanishen Tagfalter. viii + 1141 p.

SMART, P. 1975. The illustrated encyclopedia of the butterfly world. Salamander
Books Ltd., London.

Stott, C. 1782. De uitlandsche kapellen door den Heer P. Cramer.

-. 1791. In Cramer Aanbangs werk uitlandsche kappellen (1782).

Journal of The Lepidopterists’ Society
33(2), 1979, 123

BOOK REVIEW

ABERRATIONS OF BriTISH BUTTERFLIES, by A. D. A. Russwurm, 1978. E. W. Classey,
Ltd., Park Road, Faringdon, Oxon., England SN7 7DR. 151 pp., including 40
plates. £12.50 (about $25.00 U.S.).

This is basically a non-technical book of illustrations with annotations. The first
61 pages consist of introductory notes and descriptions of the specimens that are
figured in the plates. There are in excess of 300 individual specimens illustrated.
Physically the book is well produced with a pleasing format—up to Classey’s usual
high standards.

Various types of aberrations are discussed in the Introduction, but their origins
(genetic, environmental, etc.) are not. The book is basically a “picture book” of
selected aberrant butterflies, without scientific basis. The author states, “Deformed
or misshapen specimens and other abnormalities . . . are not attractive to the eye
and spoil the appearance of any cabinet drawer or coloured plate. They are not
illustrated here and we must assume that they belong to a more scientific approach
than is claimed by this book.”

The author is a fine illustrator and his watercolor renditions of the specimens
are well executed and naturally colored. Some specimens are spread in museum
fashion, while others are portrayed in natural poses. The spread specimens are
shadowed along the right margins, as if being viewed in light coming from the
upper left side of the page. I found this feature annoying as it tended to attract
the eye away from the specimens.

Although they have no standing with the International Code of Zoological Nomen-
clature, the author has supplied a “scientific” name and authority for each specimen,
with a few exceptions. The exceptions are the use of “ab. nov.” without further
comment. Of interest is an aberration of Vanessa cardui, varini Meilhan, that is
different from the usual North American aberration, elymi Rambur. I have in my
collection a cardui aberration from southern New Mexico that is intermediate
between these two forms.

One does question the reason behind producing a book such as this. It is more
of an art book than a scientific book. Perhaps it can be justified on the basis that
British lepidopterists have exhausted the more usual taxonomic frontiers. This work
will probably have some appeal in England and in Europe, where collectors seem
to be more interested in butterfly aberrations than do American collectors. I doubt
that it will have much appeal in the U.S. because of the price and the limited
geographic subject area.

CuirrorD D. Ferris, P.O. Box 3351 University Station, Laramie, Wyoming 82071.

Journal of the Lepidopterists’ Society
33(2), 1979, 124-128

REVIEW OF THE MEXICAN POLYTHRIX WATSON 1893
(HESPERIIDAE)

/

H. A. FREEMAN

1605 Lewis Drive, Garland, Texas 75041

ABSTRACT. _ Six species of Polythrix are listed from Mexico, with a key to their
identification, their synonymy, type locality, general distribution, Mexican distribu-
tion, and a plate showing the left valve of each species’ male genitalia. One name,
Polythrix alciphron (Godman & Salvin) 1893, is placed in synonymy because that
name represents a female form only.

There has been some confusion concerning members of the genus
Polythrix, particularly in the Mexican area. During the past twelve years
I have attempted to clarify the exact generic position and status of
members of this genus and their distribution in Mexico and other areas
(Freeman, 1967, 1969, 1977). In this article I present a key to the identifi-
cation of the six species which occur in Mexico. I also list the synonymy
of these species, the distribution of each based on specimens in my collec-
tion, a brief species description, and a plate showing the left valva of the
male genitalia of each species, thus simplifying determination of each
SDECIES.

Key to the Mexican Polythrix

la. No costal fold present

Ib. Costal fold present ee 3

2. Three apical spots in line; ground color light brown; lower surface of
secondaries with dark bands and spots in males, females usually have a
broad discal white area; tails in males fairly short (5 mm), in females long
(12 mm); discal bands and spots are present on the upper surface of the

secondaries but are subdued; head and thorax brown above ___. octomaculata
3a. Three apical spots present 1...) 8000) ee 4
3b. Four to five apical spots present 2.0400 a eee 5)

4a. Head and thorax brown above; discal spots on primaries semi-compact; tails
fairly short in males (5 mm), and in females longer (12 mm); discal band on
lower surface of secondaries inconspicuous __...-..-------------------2ee procerus
4b. Head and thorax brown above; discal spots on primaries compact; tails short,
in females (5 mm); discal band on lower surface of secondaries dark and

welludetined 2.20 se ie teal ie Oe guatemalaensis
Z
5a. Head and thorax brown above), 020 eee 6
5b. Head and thorax green above with intermixed brown scales __________---------- a

6a. Usually 4 apical spots; discal band not compact; ground color usually dark
brownish-black; tails in males medium length (8-10 mm), females longer
(15 mm); dark spots in space 1b on primaries evenly colored __........- asine

6b. Usually 5 apical spots; discal band not compact; ground color usually light
brown in coloration; tails in males same length as females (15 mm); dark
spots in space 1b on primaries light centered _____ Yea as J er mexicanus

VoLUME 33, NUMBER 2 125

7. Four apical spots present; rarely is there a spot in space Ib; tails short in
males (5 mm), longer in females (12-16 mm); on lower surface of second-
aries space lb from base to middle deeply grooved and at the distal end of the
eraave there is a smallverect hair tuft’on, vein Ib “2. caunus

Polythrix octomaculata (Sepp) 1848

Synonymy. decurata (H.-S) 1869; calenus (Mabille) 1888; elegans Hayward
1933; alciphron (G. & S.) 1898.

Type locality. Surinam.

General distribution. Texas to Argentina.

Mexican distribution. 6 miles south of Ciudad Valles, S. L. P. (Hotel Covadonga),
June 1969, ¢ and 2 @2 (leg. H. A. Freeman); Comala, Colima, 21 Mar. 1967,
6 (leg. Robert Wind); Salada, Colima, October 1967, 2 ¢ (leg. R. Wind);
Acahuizotla, Gro., Oct. 1950, ¢ and Oct. 1955, ¢ (leg. T. Escalante); Dos Amatos,
Veracruz, 15 Sept. 1972, @ (leg. Ramirez); Catemaco, Veracruz, 18 June 1968, 4
(leg. H. A. Freeman); Candelaria Loxicha, Oaxaca, March to November, 1968-71,
9 ¢ and @ (leg. E. C. Welling); X-Can, Quintana Roo, May 1967, 2 6 (leg. E. C.
Welling); and Muste, Chiapas, 2 Aug. 1968, ¢ and 30 Oct. 1968, ¢ (leg. E. C.
Welling collector).

Remarks. This species has three apical spots of sub-equal size forming a straight
line. The discal band is not compact. The spot in space 2 is somewhat square and
is situated midway under the larger cell spot. There is a costal spot directly over
the cell spot. The spot in space 3 is small and is located outward from the spot in
space 2 and the cell spot. There is a faint indication of the discal band on the upper
surface of the secondaries; on the lower surface the band is much darker and well
defined. The tails are fairly short in the males. There is no indication of a costal
fold which Evans (1952) says is characteristic of the genus. Head and thorax are
brown above. The female upperside is very similar to the male except the tails are
longer. On the lower surface of the secondaries specimens vary from those like
males (not having any white) to specimens which represent the type of alciphron
(Godman & Salvin) in having a large white discal area. Most of the specimens I have
examined from Mexico have some white on the lower surface of the secondaries
thus indicating that alciphron is a female form of octomaculata and thus a synonym
of octomaculata (New Synonymy ).

Polythrix asine (Hewitson) 1867

Type locality. Nicaragua.

General distribution. Mexico to Peru.

Mexican distribution. 6 miles south Ciudad Valles, S. L. P. (Hotel Covadonga),
June-July 1966-72, 28 ¢ and 14 @ (leg. H. A. Freeman); Salada, Colima, May to
June 1967, 2 ¢ (leg. Robert Wind); Comala, Colima, April 1967, ¢ (leg. R.
Wind); Presidio, Veracruz, July 1951, ¢ and @ (leg. Ramirez); Catemaco, Vera-
cruz, Aug. 1965, @ (leg. Ramirez); Candelaria Loxicha, Oaxaca, June to Sept.,
1968-71, 15 $ and @ (leg. E. C. Welling); and Muste, Chiapas, June-July 1968,
5 6 (leg. E. C. Welling).

Remarks. Usually 4 apical spots and sometimes 5. The spots increase in size
from space 8 to 6. The spot in space 9 is small and if there is a spot in space 5 it
also is small. The apical spots form a straight line. The discal band is not compact.
The spot in space 2 is fairly small and is usually located somewhat outward from
the cell spot, however in some specimens this spot is in line with the cell spot. Spot
3 is over the outer edge of the spot in space 2, well separated from the cell spot.
There may or may not be a small costal spot over the cell spot. There are usually
two dark spots in space 1b, one under the inner edge of spot 2 and the other basad.
The discal bands are prominent on the upper surface of the secondaries; however

126 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Figs. 1-6. Male genitalia, left valvae. 1, Polythrix octomaculata (Sepp). 6 mi. S.
Ciudad Valles, S. L. P. (Hotel Covadonga), 25 June 1969 (H. A. Freeman; H. A. F.);
2, Polythrix asine (Hewitson). 6 mi. S. Ciudad Valles, S. L. P. (Hotel Covadonga),
10 June 1966 (H. A. Freeman; H. A. F.); 3, Polythrix mexicanus Freeman. Paratype,
6 mi. S. Ciudad Valles, S. L. P. (Hotel Covadonga), 29 July 1966 (H. A. Freeman;
H. A. F.); 4, Polythrix procerus (Plotz). 6 mi. S. Ciudad Valles, S. L. P. (Hotel
Covadonga), 11 June 1967 (H. A. Freeman; H. A. F.); 5, Polythrix guatemalaensis
Freeman. Holotype, Sayaaxche, El Petan, Guatemala, 23 August 1963 (E. C.
Welling; A. M. N. H.); 6, Polythrix caunus (H-S.). 6 mi. S. Ciudad Valles, S. L. P.
(Hotel Covadonga), 10 June 1967 (H. A. Freeman; H. A. F.).

they are usually rather narrow. The tails are usually of medium length; however some
specimens have fairly long tails. Costal fold is well developed. Discal and apical
spots are white. The general ground coloration is dark brownish-black. Head and
thorax above brown.

VOLUME 33, NUMBER 2 1

Polythrix mexicanus Freeman 1969

Type locality. Hotel Covadonga, Ciudad Valles, S. L. P., Mexico.

General distribution. Texas to southern Mexico.

Mexican distribution. 6 miles south Ciudad Valles, S. L. P. (Hotel Covadonga),
June—Aug. 1966-72, 31 ¢ and 13 @ (leg. H. A. Freeman); Ajijic, Jalisco, 22 Oct.
1965, @ (leg. Robert Wind); Catemaco, Veracruz, June 1966-68, 4 @ (leg. H. A.
Freeman); Juchitan, Oaxaca, 17 Aug. 1964, @ (leg. H. A. Freeman); Candelaria
Loxicha, Oaxaca, Sept.—Oct. 1968-71, 3 ¢@ (leg. E. C. Welling).

Remarks. Usually 5 apical spots with the spots increasing in size from 8 to 6.
The spot in space 9 is small as is the one in space 5, which may sometimes be absent.
The discal band is not compact. The spot in space 2 is usually midway under the cell
spot. There is a small costal spot directly over the cell spot. The spot in space 3 is
about half the size of the spot in space 2 and is located over the outer edge of that
spot and outward from the cell spot. There are two dark spots in space 1b which are
light centered; one is located under the inner edge of the spot in space 2 and the
other is basad. The discal bands are well developed above and below on the
secondaries and are broader than in asine. The tails are long in comparison to any
other species of Polythrix, particularly in the males. The ground color is light brown.
The head and thorax above are light brown.

Polythrix procerus (Plotz) 1881

Synonymy. delius (Plotz) 1881; auginulus (Godman & Salvin) 1893.

Type locality. Para, Brazil.

General distribution. Mexico to Venezuela.

Mexican distribution. 6 miles south of Ciudad Valles, S. L. P. (Hotel Covadonga),
June 1966-69, 13 ¢ and 49 (leg. H. A. Freeman); Catemaco, Veracruz (Playa
Azul), 12 Aug. 1967, ¢ (leg. H. A. Freeman); and X-Can, Quintana Roo, 2 Aug.
1962, ¢ (leg. E. C. Welling).

Remarks. There are 3 apical spots in this species; the one in space 7 is small and
situated slightly inward from the ones in spaces 6 and 8. The discal spots are semi-
compact and are sordid white. The spot in space 2 is well developed and is in line
with the cell spot. In all my specimens except the male from X-Can, Quintana Roo,
there is a small spot in space 1b situated at the outer edge of spot 2. The spot in
space 3 is small and in line with the outer edge of the spot in space 2. There is only
the slightest indication of a discal band on the upper surface of the secondaries and
on the lower surface the discal band is present but inconspicuous. The costal fold is
well developed. Head and thorax above brown.

Polythrix guatemalaensis Freeman 1977

Type locality. Sayaaxche, E] Petan, Guatemala.

General distribution. Southern Mexico to Guatemala.

Mexican distribution. X-Can, Quintana Roo, 26 July 1962, 2 (allotype), (leg.
E. C. Welling).

Remarks. There are 3 apical spots in this species; the one in space 7 is smaller
than the ones in spaces 6 and 8 but in line. The spots that form the discal band are
sordid white and they are compact with a well developed spot in space 1b, a some-
what square spot in space 2, an elongated cell spot, a costal spot of about the same
width as the cell spot, and a triangular spot in space 3 which is against the spot in
space 2 and the cell spot. There is only the slightest indication of a discal band on
the upper surface of the secondaries; however on the lower surface this band is dark
and well developed. Costal fold is well developed. The tails are comparatively short
in this species. The head and thorax above brown.

128 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Polythrix caunus (Herrich-Schaffer) 1869

Synonymy. lindora (Butler) 1870.

Type locality. Unknown.

General distribution. Mexico to Paraguay.

Mexican distribution. 6 miles south of Ciudad Valles, S. L. P. (Hotel Cova-
donga), June 1966-72, 47 6 and 23 @ (leg. H. A. Freeman); and Candelaria
Loxicha, Oaxaca, 21 Sept. 1971, ¢, and 20 Aug. 1971, @ (leg. E. C. Welling).

Remarks. This species has 4 apical spots that are sub-equal. The spot in space 9 is
small and the ones in spaces 8 to 6 are progressively larger forming a straight line.
Rarely is there a small spot in space 1b under the spot in space 2. The spot in space
2 is quadrate and is located with its inner surface directly under the center of the
cell spot. The spot in space 3 is located over the outer edge of the spot in space 2
and not touching the cell spot. There is no indication of discal spots on the upper
surface of the secondaries and only slightly indicated on the lower surface. Some-
times there is a whitish spot at the lower end of the indistinct discal band in space
lc. On the lower surface of the secondaries space 1b from base to middle deeply
grooved and at the distal end of the groove there is a small erect hair tuft on vein
lb. Tails fairly short in the males and long in the females. Maculation white. Head
and thorax above green, intermixed with brown scales. Specimens in the Valles area
are slightly less green than those from Oaxaca.

ACKNOWLEDGMENTS

I thank the National Geographic Society for grants supporting this
work.

LITERATURE CITED

Evans, W. H. 1952. A Catalogue of the American Hesperiidae. Part II, Pyrginae,
Sec. 1. British Museum, London. 170 pp., pl. 10-25.

FREEMAN, H. A. 1967. Polythrix octomaculata not procerus, in Texas. J. Lepid.
S1OXCe, PAILB PATS

. 1969. Records, New Species and a New Genus of Hesperiidae from

Mexico. J. Lepid. Soc., 23 Suppl. 2. 62 pp.

. 1977. Six New Species of Hesperiidae from Mexico. J. Lepid. Soc.

31: 89-99.

Journal of the Lepidopterists’ Society
33(2), 1979, 129-134

PREDATORY BEHAVIOR IN LITHOPHANE QUERQUERA
AND OTHER SPRING CATERPILLARS

DALE F. SCHWEITZER

Dept. of Entomology, Peabody Museum, Yale University,
New Haven, Connecticut 06520

ABSTRACT. Predatory tendencies are widespread among relatively polyphagous
noctuid larvae feeding on spring foliage of forest trees but not among their more
host-specific relatives. Prey capture behavior of Lithophane querquera is complex
and highly stereotyped. Predation may be especially important during defoliator
outbreaks since larvae can change from eating foliage to eating defoliators.

It is well known among lepidopterists that certain larvae will engage
in cannibalism, especially under crowded laboratory conditions or if de-
prived of their normal food. Some noctuid species are reportedly largely
predatory in nature, e.g., Cosmia and Enargia spp. (Forbes, 1954). How-
ever, little consideration has been given to the ecological significance of
facultative larval predation and I am aware of no published accounts of
specialized predatory behavior patterns in the Noctuidae or in other
facultatively predacious Lepidoptera.

I have reared many thousands of noctuid larvae and although studying
cannibalism was not a primary purpose of such rearings, some relevant
observations were made and I suggest some ecological implications of
these predatory tendencies.

Among the Lithophanini (Noctuidae: Cuculliinae) predatory ten-
dencies are quite widespread in certain genera, but nearly absent in
others. Species showing no predatory (i.e., cannibalistic ) tendencies even
in crowded, confined, laboratory conditions included: five species of
Metaxaglaea (two presently undescribed), Chaetaglaea sericea (Mor-
rison), C. tremula (Harvey), Epiglaea decliva Grote, E. apiata Grote,
Eucirroedia pampina (Guenée), Pyreferra pettiti (Grote), P. hesperidago
(Guenée ), P. citrombra Franclemont, Lithophane semiusta (Grote), L.
patefacta (Walker) (from Wisconsin), L. signosa (Walker), Eupsilia
morrisoni (Grote) and Homoglaea hircina Morrison. Species showing
slight to moderate predatory tendencies, at least when crowded, included:
Eupsilia sidus (Guenée) (only when starving, larvae observed in sleeves
only), E. species near cirripalea (late last instar only, including when
sleeved), E. vinulenta (Grote) (late last instar only), Xylena curvi-
macula (Morrison), Lithophane bethunei (Grote and Robinson) (espe-
cially third and fourth instars), L. innominata (J. B. Smith), L. hemina
(Grote) (seldom if ever when sleeved), L. petulca (Grote), L. grotei
(Riley) (especially third and fourth instars, including sleeved larvae),

130 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Sericaglaea signata (French) (last instar only, not when sleeved) and
Jodia rufago Hiibner (rarely, only if food was wilting). Species showing
extreme predatory tendencies (i., it was rarely possible to rear more
than one per container) were: Lithophane baileyi (Grote), L. tepida
atincta (J. B. Smith), L. querquera (Grote) (sleeved larvae also highly
predacious ), and a fourth Lithophane species that ranges from southern
New Jersey southward and has generally been included with L. patefacta
in collections and by Forbes (1954).

All of the observations reported above were in crowded, laboratory
conditions except where noted. The cuttings used as food cannot be
considered optimal (Schweitzer, 1977) even though they were very
seldom visibly wilted. Sleeved larvae had natural quality food but were
crowded, about 25-50 per 51 X 82 cm sleeve. However, at least under
stress conditions, those species indicated as predatory could be expected
to eat other caterpillars in the field. Except for Eupsilia sidus and prob-
ably Jodia rufago, all of them ate other larvae even when suitable foliage
was available. Furthermore, L. bethunei is known to be a predator on
Malacosoma pupae in the field (Sanders and Dustan, 1919). The
Eurasian Eupsilia transversa (Linnaeus) is also reportedly predacious in
the field (Stokoe and Stovin, 1948). Sleeved larvae of all species of
Metaxaglaea, Chaetaglaea, Pyreferra and Homoglaea were found to be
non-predacious even after 24-48 hours of starvation.

Further information on larvae of most of the above species can be
found elsewhere (Forbes, 1954; Schweitzer, 1974, 1977). Exceptions are
Lithophane signosa which feeds only on Platanus occidentalis and one
of the undescribed Metaxaglaea which accepts and grows (but not well)
on a variety of woody plants (its natural host is unknown). Table 1
summarizes the feeding habits of predatory and non-predatory species.

Of the highly predatory species only L. querquera has been studied in
detail. I have reared nine broods, all highly cannibalistic. The last
instar larvae seem to prefer caterpillars but accept most deciduous tree
leaves and also various rosaceous fruits and flowers. The frequency of
predatory behavior, even at low densities, as well as the stereotyped
behavior described below strongly indicates this species is at least sub-
stantially predacious under natural conditions.

Based on observations of 25-30 L. querquera (two broods), larvae
exhibited the following behavior toward prey (various noctuid or decap-
itated Tenebrio larvae). When the prey approaches the larva, or is
dropped near it, the larva raises its anterior portion slightly and begins
waving to each side, usually rather slowly. When contact is made, the
prey is grasped with the true legs and the attacker works quickly to the
caudal region where the initial bite is made (Fig. 1), except in the case

VoLUME 33, NUMBER 2 1i3ul

TaBLE 1. Larval feeding patterns of predacious and non-predacious Lithophanini.
Polyphagous species are those feeding regularly on two or more plant families,
facultatively polyphagous species are those appearing to have definite food preferences
and restricted feeders can complete development only on a limited array of plants
(see Schweitzer, 1977).

Feeding pattern

Predatory Facultatively
tendency Genus Polyphagous polyphagous Restricted Uncertain
None Lithophane 0 2 3!
Pyreferra — - 3
Eupsilia 1 0 —
Homoglaea = - I
Metaxaglaea 1 0 3 I)
Epiglaea - if if
Chaetaglaea = 2 -
Eucirroedia = = I
total non-predacious 2 5 9 2
Slight to
moderate Xylena 1 0 0
Lithophane 4 if 0
Eupsilia 2 if —
Sericaglaea I - - -
Jodia — = 1,
Extreme Lithophane 3 iP 0
total predacious Wal 3 1 0

_ Note: The symbol — indicates no species in that genus exhibits the indicated feeding strategy
in the eastern United States; 0 indicates that no species exhibiting a particular feeding pattern
was found to fit into the predation category indicated.

of decapitated Tenebrio in which case feeding usually starts at the
wound. Frequently the entire prey is eaten, but rather often the head
capsule is discarded. Prey may vary considerably in size range and can
be larger than the attacker. Occasionally L. querquera larvae encounter
non-moving prey and bite into the caudal portion quickly, omitting the
waving motions. The advantage of grasping the prey caudally may be
that this prevents it from jumping or dropping away. Both L. querquera
and L. t. atincta larvae have also been observed to turn and run down
moving caterpillars.

Table 2 presents data from an experiment designed to determine if
L. querquera larvae could grow well on an exclusively vegetable diet.
All larvae hatched 16 May 1975 and were reared individually in 230 ml
glass jars. Food was primarily foliage of Pyrus <purpurea for the first
five instars. Thereafter, half of the larvae were maintained on this diet,
with fresh food daily, during the last instar. The others were also given
noctuid or decapitated Tenebrio larvae about every second day. These
prey larvae were always eaten, and were often taken within 4 or 5

132 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fig. 1. Last instar larva of Lithophane querquera (Southford, New Haven Co.,
CT.) beginning to eat a third instar Xylena curvimacula (Sunderland, Franklin Co.,
MA.) (Photo by William Sacco, Peabody Museum Photography Laboratory: twice
life size).

seconds. In many cases small amounts of foliage were subsequently
eaten. The final body weights are very close for both groups and neither
survival nor growth rates (based on maturation date) differ statistically.
Thus L. querquera larvae are clearly not obligately predatory. It is
somewhat surprising that no advantage can be shown for the predators.
Perhaps the Tenebrio larvae which constituted over half of the prey are
a poor food, or the Pyrus may be exceptionally suitable.

Predatory behavior probably serves to reduce the effects of competition

TABLE 2. Comparisons of body weights for sibling L. querquera larvae fed
foliage vs. mixed foliage-insect diets during their last instar. Weights were taken
the afternoon following the last night of feeding and are in grams.

Food in last instar

Date matured Foliage (N = 16) Insects and foliage (N = 17)
16 June .2818
17 June 0203
18 June .2856
19 June PASAT .2842
21 June 2813 2822
22 June .2260
23 June PAO .2614, .2377
26 June PALLIUS) .1952, .2801, .2480
27 June 0068, 200)
28 June 2302.) 2000 .2610
29 June .2631
3 July PRBS:
mean weight .2658 .2617
% survival 56.3 716.5
Comparison of survival data gives x? = 1.58, p >> .05. Comparison of maturation dates by a

Mann-Whitney U-test (Siegel, 1956; 2 tailed large sample procedure) gives U = 39.5, p = .1936.

VoLUME 33, NUMBER 2 ILI)

for food from other caterpillars in certain situations. Even before the
disturbance brought on by European man and the pests he has imported
to North America, occasional spring canopy defoliation probably occurred
in the eastern deciduous forests. Likely defoliators would include the
several Geometridae commonly known as canker worms (Craighead,
1950). In fact, Lithophane larvae themselves occasionally cause local
canopy defoliation (Craighead, 1950; Rings, 1968, 1973). A facultative
predator (or cannibal) could switch from eating foliage to eating defoli-
ators during outbreak periods. It is not known whether any lithophanine
larvae will eat gypsy moth (Lymantria dispar | Linnaeus] ) larvae, but us-
ually defoliations caused by that species occur after most Lithophanini
have finished feeding. It is interesting to note that one of two generalized
feeders (Table 1) listed as nonpredatory, Eupsilia morrisoni, apparently
feeds on understory plants in its late instar (Rings, 1969) and thus would
be little affected by canopy defoliation.

Apparently, however, it is difficult for specialized feeding larvae to
evolve (or retain) predatory behavior since nine out of ten restricted
feeders (and most others with distinct preferences) are non-predatory
(Table 1). In addition, I find no evidence of cannibalism among ten
early feeding species of Catocala I have reared. Larvae of this genus
are highly specialized feeders and none is reported as cannibalistic even
in confinement (Sargent, 1976). Furthermore, the clear lack of can-
nibalism among crowded Lithophane signosa and less crowded L.
semiusta and L. patefacta (whose very near relatives include predacious
species ) suggests this tendency may be lost as specialized feeding habits
evolve, assuming that these species evolved from more generalized rela-
tives. This assumption is questionable for L. semiusta but seems very
likely for the others.

At the other extreme L. querquera represents a generalist that has
evolved into a substantially predacious niche. In addition to the be-
haviors already described, it seems to differ from other Lithophanini by
being somewhat more active, both nocturnally and diurnally. Presum-
ably, increased locomotor activity increases the chance of encountering
suitable prey.

Another interesting feature of L. querquera that perhaps related to
its hyperactivity is its unusual coloration. This species may be warningly
or mimetically colored since, unlike all other known Lithophane larvae,
L. querquera larvae do not appear to be cryptic. They are greyish or
bluish to whitish with a bright yellow pattern (Schweitzer, 1974, 1977).
The color and pattern, however, are quite close to those of Pyreferra
ceromatica, hesperidago, and citrombra. These three feed almost entirely
on Hamamelis virginiana in southern New England (Schweitzer, 1977),

134 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

where they are easily found on the undersides of the leaves in May and
June.t Possibly they derive some toxic or noxious substance from this
plant and are mimicked by the less numerous, but presumably edible,
L. querquera larvae. The possibilities of mimicry or unpalatability could
be investigated experimentally if stock of these species were available.
The predatory habits of noctuid larvae have received very little at-
tention from ecologists or entomologists. A more precise system of
classifying predatory tendencies could be devised. The observations
presented here suggest that predatory tendencies may be an important
factor favoring non-restricted feeding habits among spring canopy feed-
ing noctuid larvae. It seems likely that the impact of severe competition
for food would be less for such larvae than for their restricted, non-
predatory relatives. Experimental studies of predation during simulated
cankerworm outbreaks will be reported elsewhere. In some instances
cannibalism may be an important mechanism of self-regulation.

LITERATURE CITED

CRrAIGHEAD, F. C. 1950. Insect Enemies of Eastern Forests. U.S. Dept. Agric.
Misc. Publ. 679 pp.

Forses, W. T. M. 1954. Lepidoptera of New York and Neighboring States. III.
Noctuidae. Cornell Univ. Agric. Exp. Sta. Mem. 329. 433 pp.

Rincs, R. W. 1968. Contributions to the bionomics of the green fruitworms: the
life history of Lithophane laticinerea. J. Econ. Entomol. 62: 1388-1393.

. 1969. Larval Behavior of Eupsilia morrisoni. Ann. Entomol. Soc. Amer.

62: 544-549.

1973. Contribution to the bionomics of green fruitworms: the life history
of Lithophane antennata. J. Econ. Entomol. 66; 364-368.

SANDERS, G. E. & A. G. Dusran. 1919. The Fruitworms of the Apple in Nova
Scotia. Can. Dept. Agric. Tech. Bull. 17: 5-28.

SARGENT, T. D. 1976. Legion of Night: The Underwing Moths. Univ. Massachu-
setts Press. Amherst, Mass. 222 pp.

SCHWEITZER, D. F. 1974. Notes on the biology and distribution of the Cuculliinae
(Noctuidae). J. Lepid. Soc. 28: 5-21.

1977. Lite History Strategies of the Lithophanini (Lepidoptera: Noctuidae,
Cuculliinae), the Winter Moths. Ph. D. thesis, University of Massachusetts, Dept.
of Zoology. 304 pp.

SieGAL, S. 1956. Nonparametric Statistics for the Behavioral Sciences. McGraw-
Hill Book Co. New York. 312 pp.

STOKOE, W. J. & G. H. T. Srovin. 1948. The Caterpillars of British Moths, ser. 1.
F’. Warne & Co., London and New York. 407 pp.

1P. citrombra is nepem cy a Corylus feeder (Forbes, 1954), but southern New England larvae
accept only Hamamelis (2 spp.) and I have collected them on H. virginiana in Massachusetts
and at Philadelphia, Pennsylvania. P. ceromatica has apparently not been collected in New
England or neighboring regions for about 60 years, but was once fairly common there.

Journal of the Lepidopterists’ Society
33(2), 1979, 135-136

A HOLOTYPE DESIGNATION FOR PAPILIO CINYRAS RIDENS
MASTERS 1971 (PAPILIONIDAE)

JoHn H. Masters

25711 N. Vista Fairways Drive, Valencia, California 91355

ABSTRACT. In an earlier paper (Masters, 1971) the name ridens was elevated
to subspecific rank for the first time. Although the name was attributed to Fassl,
the International Code of Zoological Nomenclature provides that if an infrasub-
specific name is elevated to the rank of the species group, it will take the date and
authorship of its elevation. Thus Papilio cinyras ridens should be attributed to Masters
(1971). A holotype male is designated and will be deposited in the Los Angeles
County Museum of Natural History, Los Angeles. The specimen is from Buenavista,
Dept. Santa Cruz, Bolivia and was collected by Franz Steinbach in 1962.

In an earlier paper ( Masters, 1971), I elevated an old infrasubspecific
name of Fass] (1915) to subspecific status as Papilio cinyras ridens at-
tributing the name to Fassl. At the time I thought I was elevating and
conserving an old name, however the provisions of the International Code
of Zoological Nomenclature (Anonymous, 1964) provide (Article 10b)
that “A name first established with infrasubspecific rank becomes avail-
able if the taxon in question is elevated to a rank of the species-group,
and takes the date and authorship of its elevation.” Thus the name ridens
should be attributed to Masters with the authorship date of 1971. There
is an obvious problem here; at the time of publication I did not realize
that I was establishing a new taxon, and while identification characters
and geographic criteria were given, no type specimens were established.
In order to rectify this situation, I hereby establish the following types:

Papilio cinyras ridens Masters 1971

Holotype. Male, Buenavista, Dept. Santa Cruz, Bolivia, collected by Franz Stein-
bach (May 1962). Deposited in the Los Angeles County Museum of Natural History,
Los Angeles.

Paratypes. 19 males, 3 females, same data as holotype (various dates). Presently
in the author’s collection but will be distributed (upon request) to any persons
working on taxonomy of Neotropic Papilionidae.

Pipilio cinyras ridens is well-depicted in color by Fass] (1915). My
original paper ( Masters, 1971) provides a sketch showing the distinguish-
ing characters between it and other subspecies of Papilio cinyras; it also
gives the criteria for separating Papilio cinyras from Papilio thoas (all of
which were combined under Papilio thoas by Rothschild and Jordan,
1906). I regard Papilio cinyras to comprise the following:

(a) P. c. cinyras Menetries, Peru

136 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

(b) P. c. ridens Masters, Eastern Bolivia
(c) P. c. brasiliensis Rothschild & Jordan, Brazil and southward.

LITERATURE CITED

AnonyMous. 1964. International Code of Zoological Nomenclature adopted by
the XV International Congress of Zoology. Edited by N. R. Stoll, R. Ph. Dollfus,
J. Forest, N. D. Riley, C. W. Sabrosky, C. W. Wright and R. V. Melville.
International Trust for Zoological Nomenclature, London.

Fass, A. H. 1915. Neue Schmetterlingsformen aus Sud-Amerika Papilioniden.
Deutsche Entomol. Zeitschrift Irus 29: 186-189.

Masters, J. H. 1971. Papilio cinyras ridens Fassl: a new status (Lepidoptera:
Papilionidae). Entomol. Rec. 83: 83-86.

RotrHscHitp, W. & K. JorpAn. 1906. A revision of the American Papilios. Novit.
Zool. 13: 27-752.

Journal of the Lepidopterists’ Society
33(2), 1979, 136

MELIPOTIS INDOMITA (NOCTUIDAE) IN HAWAII AND CONNECTICUT

Recently while curating material of Melipotis indomita (Walker) at the Peabody
Museum, Yale University, I encountered two specimens from unusual localities
which seemed to merit ‘a brief note. The data are as follows:

HAWAII: Oahu, Honolulu Co., Honolulu, 28 May 1974 leg. Joseph G. Gall, a
fresh male.

CONNECTICUT: Litchfield Co., Washington, at home lights, 31 July 1958, leg.
Sidney A. Hessel, a worn female.

Forbes (1954, Lepidoptera of New York and Neighboring States, Part 3, Cornell
Agric. Exp. Sta. Mem. 329) gives the distribution as, “Southern states; reported
northward at Kittery Point, Maine, Long Island, St. Louis, Missouri (not rare) and
Delaware.” Most Peabody Museum specimens are from Arizona.

The Connecticut record is remarkable since the specimen was taken in the Litch-
field hills, about 47 km from the coast. It probably arrived there by natural dispersal.
The species probably has been accidentally introduced to Hawaii by human trans-
port. No species of Melipotis Hiibner or the closely related Drasteria Hiibner (sensu
Forbes, 1954) is reported from the Hawaiian Islands by Zimmerman (1958, Insects
of Hawaii, vol. 7, Univ. of Hawaii Press, Honolulu).

Date F. ScHWEITzER, Curatorial Associate, Entomology, Peabody Museum, Yale
University, New Haven, Connecticut 06520.

Journal of the Lepidopterists’ Society
33(2), 1979, 137-138

THE TYPE LOCALITY OF ARGYNNIS ZERENE BOISDUVAL
(NYMPHALIDAE): A CORRECTION

Joun H. Masters

25711 N. Vista Fairways Drive, Valencia, California 91355

ABSTRACT. Argynnis zerene Boisduval was described from material collected
by P. J. M. Lorquin during 1850 and/or 1851 in California. The original description
gives the locality as the low mountains of California. Dos Passos and Grey re-
stricted the type locality to Yosemite Valley, Mariposa County, California. This is not
possible, since Yosemite Valley was not discovered until 1851 and Lorquin could
not have collected there before 1856—four years after the published description of
zerene. The type locality is corrected and redesignated as vicinity of Agua Fria,
Mariposa County, California. This taxon is now subjectively placed in the genus
Speyeria.

Argynnis zerene (a species now subjectively placed in the genus
Speyeria) was described by Jean Boisduval (1852) from material col-
lected in California in 1850 and/or 1851 by P. J. M. Lorquin. In the
original description, Boisduval stated simply that zerene occurred in the
low mountains of California in June. Dos Passos and Grey (1947), with-
out citing any reasons, restricted the type locality to Yosemite Valley,
Mariposa County, California.

This is impossible. The discovery of Yosemite Valley was on 27 March
1851, by a military party under the command of Major J. D. Savage,
during the Mariposa Indian War (Farquhar, 1965: 72). Yosemite was
not visited again until late June 1855 when J. M. Hutchings led a party
of four there. Hutchings is known to have collected butterflies for Henry
Edwards at a later time (Edwards, 1878), but certainly did not collect
any at this time. The Hutchings Expedition brought publicity to Yosemite
and by the close of 1855, total tourist travel to Yosemite had reached 42
(Farquhar 1965: 117-118). The first structure in Yosemite was erected
in 1857 with the intent to operate it as a hotel, however it was 1859 before
any appreciable travel to Yosemite took place (Farquhar, 1965). If
Lorquin ever collected in Yosemite, it is impossible for him to have done
so before 1856 and extremely unlikely before 1859.

The exact localities where Lorquin collected in 1850 and 1851 are not
known. We do know that he collected in the vicinity of San Francisco
and the placer mines of Tuolumne County (cf. Grinnell, 1904). In
another paper (Masters, in preparation) I have determined that it is
highly probable that he spent the spring and early summer of 1851 in an
area bounded by Mokelumne Hill (Calaveras Co.) on the north and
Coarsegold (Madera Co.) on the south. An area which includes the
mining towns of Murphys and Angel’s Camp (Calaveras Co.), Sonora

138 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

and Columbia (Tuolumne Co.), and Agua Fria (Mariposa Co.). He
could have been in any of these towns, or in several of them.

Of these possibilities, Agua Fria is the closest to Yosemite. As it is
quite possible that Lorquin could have been in Agua Fria, I hereby
designate it as the corrected type locality for Argynnis zerene. There
is nothing left of Agua Fria now, but in 1850 and ‘51 it was a booming
placer gold camp and the county seat of Mariposa County. The town was
located on Agua Fria Creek just west of the present town of Mariposa
and some 35 miles southwest of Yosemite Valley.

Dos Passos and Grey (1947) considered Argynnis monticola Behr
(types taken in Yosemite Valley in 1863 by the California Geological
Survey ) a subjective synonym of Speyeria zerene. Considering Yosemite
Valley as the type locality for S. zerene facilitated this conclusion. The
correction of the type locality to Agua Fria should not upset the taxo-
nomic conclusions of Dos Passos and Grey and will not change any of the
synonymies per contemporary usage. Agua Fria is in the same biotic
province as Yosemite Valley and it is unlikely that any future systematist
would consider populations from the two localities as separate and dis-
tinct subspecies.

The type specimen of Argynnis zerene is in the U.S. National Museum
in Washington, ex collection Barnes, ex collection Oberthur, ex collection
Boisduval.

LITERATURE CITED

BotspuvaAL, J. 1852. Lepidoptéres de la Californie. Ann. Soc. Entomol. France,
2nd ser. 10: 275-324.

Dos Passos, C. F. & L. P. Grey. 1947. Systematic catalogue of Speyeria (Lepi-
doptera, Nymphalidae) with designations of types and fixations of type localities.
Am. Mus. Novit. 1370: 1-30.

Epwarps, H. 1878. Pacific Coast Lepidoptera. No. 30. Proc. Calif. Acad. Sci.
7; 11-14.

FarQuHar, F. P. 1965. History of the Sierra Nevada. Univ. Calif. Press, Berkeley.

GRINNEL, F. 1904. An early collector in California. Entomol. News 15: 202-204.

Journal of the Lepidopterists’ Society
33(2), 1979, 139-143

NOTES ON CHILEAN OECOPHORIDAE

J. F. Gates CLARKE
Dept. of Entomology, Smithsonian Institution, Washington, D.C. 20560

ABSTRACT. The rediscovery of Hyperskeles choreutidea Butler is recorded and
the genitalia are illustrated for the first time. Tyriomorpha Meyrick, 1918 (= Mattea
Duckworth, 1966), New Synonymy, is noted. The information contained in this
short paper was not available in time to be included in my recent paper (Clarke,
1978), the latter in press over two years.

Hyperskeles Butler

Hyperskeles Butler, 1883: 78 (Type-species: Hyperskeles choreutidea Butler loc.
cit.: 79 [by monotypy].)—Fletcher, 1929: 114.—Gaede, in Bryk, 1939: 398.

The original description of this genus is as follows: “Allied to
Oecophora, aspect of Dasycera; secondaries broader than in either genus;
the primaries rounded at apex, but with the external angle well defined
and consequently with short fringe; antennae filiform; palpi slender,
porrect, long and widely separated; legs long and thick, but not fringed.”

An emended generic description follows:

Labial palpus smooth, rather slender, recurved, slightly exceeding vertex; third
segment nearly as long as second, acute. Tongue well developed, thickly scaled;
maxillary palpus minute, slender, free. Head with closely appressed scales,
small sidetufts spreading; ocellus absent. Antenna filiform; scape without pecten.
Thorax smooth. Forewing smooth, costa slightly arched, termen convex, 11 veins;
1b furcate; 1c preserved at margin; 2 and 3 connate from angle of cell; 4 approximate
to 3; 5 nearer to 4 than to 6; 6 to termen slightly below apex; 7 and 8 coincident to
costa slightly before apex; 9 nearer to 7 + 8 than to 10; 11 from middle. Hindwing
with 8 veins; 2 remote from 3; 3 and 4 stalked; 4 to 7 parallel and equidistant. Hind
tibia smooth. Abdominal terga weakly setose; setae deciduous.

Male genitalia with uncus and gnathos well-developed, socius absent. Vesica
armed.

Female genitalia without signum.

This genus keys to Despina in my key (Clarke, 1978:3) but differs from that
genus by the filiform antenna, 2 and 3 connate, and the broadly rounded termen in
forewing.

Hyperskeles choreutidea Butler
Figs. 1-5

Hyperskeles choreutidea Butler, 1883: 78.—Bartlett-Calvert, 1886: 346.
Hyperskeles choreutidia [sic] Gaede, in Bryk, 1939: 398.

Male genitalia slide USNM 77489 (AB June 18, 1934). Harpe triangular, cucullus
bluntly pointed; sacculus sclerotized, terminating in a pointed process, the latter
extending two-thirds across middle of harpe; base of harpe with lunate, setose pad
on inner surface. Gnathos a slender hook. Uncus clavate; inner surface, distad,
clothed with setae. Vinculum U-shaped. Tegumen about as long as uncus. Anellus

140 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Figs. 1-5. Hyperskeles choreutidea Butler: 1, dorsal view of adult; 2, venation
of wings; 3, ventral view of male genitalia with left harpe and aedeagus removed;
4, aedeagus; 5, ventral view of female genitalia.

an oval sclerotized plate with deep median cleft. Aedeagus slender, nearly straight,
distally pointed; vesica armed with fine cornuti.

Female genitalia slide USNM 24378. Ostium transverse, crescentic. Antrum a
sclerotized cone. Inception of ductus seminalis ventrolaterally slightly before antrum.
Ductus bursae membranous. Bursae copulatrix membranous with a slightly rugose
section posteriorly. Signum absent.

Holotype: British Museum (Natural History).

Type locality: Valdivia.

Distribution: Chile, Argentina.

The known distribution of the species, in addition to Valdivia, the type locality, is
as follows: Chile, ¢, 9, Callipulli (“a town in Chile, in the province of Malleco,
90 mi SE of Conceptién”), March, Silva; S. Chile, ¢, Los Muermos, (Forest) 19
Jan. 51. R. Michelbacher; Argentina, 9, prov. Chibut, El Tutbio. Lago Puelo,
25.111.1961, Gy Topal.

Foodplant: Unknown.

As far as I am able to ascertain, this species has been “lost” since 1856
(p. 346) when Bartlett-Calvert listed it under the Gelechiidae in his
catalogue of the Lepidoptera of Chile. Gaede listed it (1939:398) under

the Oecophoridae but Fletcher (1929:114) listed the genus and species
as questionably oecophorid.

VoLUME 33, NUMBER 2 14]

Figs. 1-5. Continued.

142 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

In the United States National Museum of Natural History there are
two specimens (é & ¢) that had remained hidden in the unworked
Choreutidae since the early 1930’s until discovered recently by John
Heppner while he was working on that family. Heppner acquired a
third specimen (2) from Dr. L. Gozmany of the Zoological Department,
Hungarian Natural History Museum, Budapest, and a fourth specimen
(2) from the Los Angeles County Museum where they were thought
to be choreutid. When Heppner brought the specimens to me, aware
that they were not choreutids, I recognized them immediately from a
colored illustration (origin unknown) in the collection of the United
States National Museum of Natural History.

Through the courtesy of Dr. Klaus Sattler, British Museum (Natural
History) I learned (in litt.) that the type 2 of choreutidea is in that
museum where it had been placed in the Gelechiidae.

The following synonymy was brought te my attention by Dr. John
Heppner and Dr. Klaus Sattler:

Tyriomorpha Meyrick, 1918, Exotic Microlepidoptera, 2: 191 (Type-species:
Cryptolechia phoenissa Butler, 1883, Trans. Ent. Soc. London, 81, pl. 11, figs.
12, 12a [by original designation]. )

Mattea Duckworth, 1966. Proc. U.S. National Museum, 119 (No. 3540): 2 (Type-
species: Cryptolechia phoenissa Butler, 1883, Trans. Ent. Soc. London, 81, pi. 11,
figs. 12, 12a [by original designation].) New Synonymy.

Cryptolechia phoenissa Butler was originally placed in the Gelechiidae,
where it remained until Duckworth (1966) placed it correctly in the
Oecophoridae. When Meyrick (1918) described Tyriomorpha he added
to the confusion by placing it in the Glyphipterygidae. Clarke (1978:
7) also placed the species correctly in the Oecophoridae, but in the genus
Mattea Duckworth.

ACKNOWLEDGMENTS

The drawings for this paper were made by Margaret Garrison and the
photograph was produced by Victor Krantz, Smithsonian Institution.

LITERATURE CITED

BartTLetr-CaLvert, W. 1886. Catalogo de los Lepiddpteros Rhopaldceros i
Heteréceros de Chile. Anales de la Universidad de Chile (Santiago) 69:
ol1—352.

CLARKE, J. F. Gates. 1978. Neotropical Microlepidoptera XXI: New Genera
and Species of Oecophoridae from Chile. Smithsonian Contributions to Zoology,
No. 273: 1-80, figs. 1-54, pl. 1-6.

Duckwortu, W. D. 1966. Neotropical Microlepidoptera X: Systematic position
of Two Taxa Erroneously placed in the Family Stenomidae (Lepidoptera).
Proc. U.S. National Museum, 119 (No. 3540): 1-6, pl. 1.

VOLUME 33, NUMBER 2 143

FLetcuer, T. B. 1929. A List of the Generic Names Used for Microlepidoptera.
Memoirs of the Department of Agriculture in India. Entomological Series,
11:i-ix, 1-244.

GaEebE, M. 1939. In Bryk, Lepidopterorum Catalogus, 92: 209-476. s’Gravenhage:
W. Junk.

Journal of The Lepidopterists’ Society
33(2), 1979, 143-145

GENERAL NOTES

NOTES OF MARYLAND LEPIDOPTERA. 6. OCCURRENCE OF
BOLORIA SELENE (NYMPHALIDAE) IN MARYLAND

In 1941, Clark (J. Wash. Acad. Sci. 31: 381-384) named a new subspecies of
Boloria selene from specimens he had caught near Beltsville, Maryland in 1929. It
was described as single brooded and “. . . resembling Brenthis selene myrina but
larger . . . and with the ground color above darker and more reddish and the black
markings broader and heavier. . . .” Clark had pictured the type earlier in his
Butterflies of the District of Columbia and Vicinity (1932, U.S. National Mus. Bull.
157) and later (Clark and Clark 1951, Smithsonian Misc. Coll. 116, No. 7. 239 p.)
reported that the 1929 specimens were the last to be found in Beltsville.

This subspecies, named marilandica (Fig. 1), reappeared in Largo, Maryland in
1941 when Dr. Warren Wagner, Jr. captured at least one specimen there (Clark
and Clark, loc. cit.). In 1948 and 1949, one of us (WAA) caught several specimens
which were first identified as myrina, but when they were shown to A. H. Clark
himself in late 1952, he identified them as typical marilandica and noted that he
was happy to know of another locality where they could be found.

Since then several more colonies have been located. They seem to be clustered
around the Fall Line as it makes its way in a northeast-southwest direction through
Maryland (Fig. 2). (The Fall Line is a line of rocky falls on the courses of the
many streams and rivers that in Maryland empty into the Chesapeake Bay. It divides
the low, flat Atlantic coastal plain from the gentle, rolling hills of the piedmont. )
- However, we know of no specimens caught later than 1966. This apparent disap-
pearance may be due either to our not collecting at the right time in the swampy
areas where the insects occur, or to their actual extinction, due to the pressures of
human population extension into their areas which is occurring rapidly in the
Baltimore-Washington-Philadelphia corridor.

An analysis of the capture dates of our “marilandica” specimens suggests there are
three broods: late May to mid-June, mid-July to early August, and mid-August to
late September.

The populations of the different areas in which we collected were somewhat
variable. Many specimens were larger and darker than normal, agreeing with the
description of “marilandica.” However, we also found a few that agreed neither with
“marilandica” or myrina, some being larger but lighter in color, others being smaller
but having very thick, heavy black markings. Likewise, there were two colonies that
existed close by those of “marilandica” in which all the specimens were small and
only a few conforming to “marilandica” could be found. One of us (RSS) col-
lected myrina earlier (2 September 1941) in Lutherville, Maryland, which is very
near Stevenson (Fig. 2), and these specimens do not conform to “marilandica.”

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

144

°
?

ins,

8. same, ventral.

2

5. female, Hark

?

1. male, Eklo, 17 July 1948

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VoLUME 33, NUMBER 2 145

79° 76° es 77° 76°
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BEESEACHIAN AS PIEDMONT

FALL ZONE

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ect
e “os at
wyeo ’

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CARR oe j@4) ae, 2D
———— — 5 Se af fh > ae
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a8 Coastal Plain sediments of gravel,

} sand, silt, and clay

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Sedimentary rocks — limestone,
sandstone, shale, and siltstone

MARYLAND

SCALE

38°
© 10 20 30 40 50 MILES
SSE ——

Fig. 2. Distribution of B. s. myrina, form “marilandica,” in Maryland. Map
adapted from Brush, Lenk & Smith (1977, Dept. Geography and Environmental
Engineering, Johns Hopkins Univ., Baltimore, Md. 81 p.) and reproduced by written
permission of the authors. 1. Largo, Prince Georges County; 2. Beltsville, Prince
Georges County; 3. Stevenson, Baltimore County; 4. Eklo, Baltimore County; 5.
Coopstown, Harford County; 6. Harkins, Harford County; and 7. Chesapeake City,
Cecil County.

In the two small colonies in Harford County the specimens were all “marilandica.”
Only one specimen was obtained from the Chesapeake City (Cecil County) locale.

These observations and data support the recent revision of selene by Kohler (1977,
J. Lepid. Soc. 31: 243-268) which places “marilandica” in synonymy with myrina,
since “marilandica” is not single-brooded as Clark had thought, and specimens of
this form are found in some populations north of Maryland (Kohler, loc. cit., Fig. 2).
In fact, the type specimen of the species figured by Cramer (1779, Papillons
exotique des trois parties due monde, |’Asie, Afrique et !’Amerique. Baalde, Am-
sterdam; Barthelmy Wild, Utrecht. 4 vol.), although not comparable with “mari-
landica” in size, appears to be just as heavily marked as “marilandica.” The type
specimen is from New York.

NEW MARYLAND RECORDS. Baltimore County: Eklo, 28 September 1947;
17 July 1948; 25 July 1948; 15 August 1948; 29 August 1948; 30 May 1949; 6 June
1949; 9 June 1949; 13 September 1962; 11 July 1963; 28 May 1964. Stevenson,
14 August 1965; 4 June 1966. Lutherville, 2 September 1941. Cecil County:
Chesapeake City, 21 August 1952. Harford County: Harkins, 2 June 1966; 14 July
1966. Coopstown, 14 July 1966. Prince Georges County: Beltsville, 21 July 1961
(topotype caught by Mr. William Field and now in the collection of the U. S.
National Museum, Washington, D.C.).

Wit1t1aM A. ANDERSEN, 220 Melanchton Avenue, Lutherville, Maryland 21093.
Rosert S. Simmons, 1305 Light Street, Baltimore, Maryland 21230.

146 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Journal of The Lepidopterists’ Society
33(2), 1979, 146-147

TWO SOUTHERN COSSIDS (COSSIDAE) IN THE NEW JERSEY
PINE BARRENS

I have been collecting Lepidoptera in the New Jersey Pine Barrens for over ten
years and have been working off and on at a regional moth list. This area is well
known for harboring many unusual species, including more or less disjunct popula-
tions of both boreal and southern coastal plain species.

However, my recent discovery of two southern cossids there came as quite a
surprise. The first of these, a single male of Givira anna (Dyar) (Fig. 1, above) was
collected at a 20W black light by me at Batsto, Burlington Co. on 21 June 1969. It
was identified at the American Museum of Natural History in June 1973 with the
help of Dr. Alexander B. Klots and later confirmed by Dr. John G. Franclemont in
August 1975. It is in my collection. Barnes and McDunnough (1911, Revision of
the Cossidae of North America, Decatur, Illinois, Review Press) had records of this
species only from Florida. Kimball (1965, The Lepidoptera of Florida, State of Fla.,
Div. of Plant Industry) gave records covering much of the state and indicated the
food to be pine. I also have a male of this species from Florence, South Carolina,
14 August 1963, leg. V. M. Kirk. The Peabody Museurn collection at Yale Uni-
versity contains at least three males from McClellanville, South Carolina, taken at
the Wedge Plantation, 4-14 August 1967, leg. Charles W. Porter.

The second species is Inguromorpha basalis (Walker) (Fig. 1, below), a male of
which was taken at Batsto on 2 July 1972 in a black light trap operated for me by
Annie Carter. The specimen is in my collection. I again encountered this species
at the same locality on 20 June 1977 when I tock a much larger male in a MV

Nig. 1. Two southern cossids taken in the New Jersey Pine Barrens: above,
Givira anna ¢, Batsto, Burlington Co., New Jersey, 21 June 1969, leg. D. F.
Schweitzer; below, Inguromorpha basalis 6, same locality, 2 July 1972, leg. A.
Carter. Both specimens life-size.

VoLUME 33, NUMBER 2 147

Robinson trap. Unfortunately the apical regions of both forewings are badly
damaged, probably from flying inside the trap. This specimen is at the Peabody
Museum, Yale University.

The 1972 specimen was confirmed by Dr. Franclemont along with the above
Givira. Barnes and McDunnough (op. cit.) record it only from Florida and Kimball’s
records (op. cit.) cover much of that state. All of these specimens were collected
within 20 m to the north of the Batsto Nature Center, situated on the top of the
small hill near the east bank of the Batsto river, just above the dam. The surrounding
vegetation includes a woodlot of various oaks and adventive species and extensive
areas of essentially natural oak-pine and pine-oak forests extending more or less
unbroken for hundreds of square kilometers, especially to the north. The pines are
Pinus echinata Mill., and P. rigida Mill., with the former predominating at the im-
mediate area of the captures.

Dare F. ScHweITzER, Curatorial Associate, Entomology, Peabody Museum, Yale
University, New Haven, Connecticut 06520.

Journal of The Lepidopterists’ Society
33(2), 1979, 147-148

A MELANISTIC SPECIMEN OF ANTHERAEA POLYPHEMUS POLYPHEMUS
(SATURNIIDAE )

On 2 June 1975, the senior author received a living specimen of Antheraea poly-
phemus polyphemus (Cramer) that was most unusual in coloration (Figs. 1-4). The
moth, a female, had eclosed on 1 June from a cocoon found approximately two
weeks previously on a fence in Winnipeg, Manitoba. The cocoon had been given

Figs. 1-4. Antheraea polyphemus (Cramer): 1. typical female from Winnipeg,
dorsum; 2. typical female, venter; 3. melanistic female, dorsum; 4. melanistic female,
venter. (Photos by W. B. Preston).

148 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

to a school student, Miss Darcy Berry, who, on realizing the unusual nature of the
moth, donated it to the Manitoba Museum of Man and Nature.

The general coloration is dark brown, but with the recognizable markings of
A. p. polyphemus. The ventral surface is of the same colour as the dorsum but is
contrastingly marked as is typical of specimens from Manitoba (Figs. 2, 4). In
wingspan the mounted specimen measures 114 mm and the greatest length of the
right front wing is 61 mm.

There are some references to melanistic A. polyphemus in the literature. Holland
(1903, The Moth Book. Doubleday, New York. 479 p.) mentioned “. . . one or two
fine melanic specimens, in which the wings are almost wholly black on the upper
side.” Packard (1914, Mem. Natl. Acad. Sci. 12: 207) referred to Holland’s speci-
mens and (p. 205) mentioned three purplish coloured specimens reared from
cocoons from Macon, Georgia. Ferguson (in Dominick, R. B. et al. 1972, The Moths
of America North of Mexico, fasc. 20.2B Bombicoidea (in part) ) considered
melanistic specimens to be very rare. The authors of the present paper are unaware
of any prior published illustration of a melanistic A. polyphemus.

We wish to thank Mr. Richard Westwood, University of Manitoba, Winnipeg,
and Mr. C. S. Quelch, Winnipeg, for examining their collections for melanistic speci-
mens.

WILLIAM B. PRESTON AND WM. BriAN McKiLuLop, Manitoba Museum of Man and
Nature, 190 Rupert Avenue, Winnipeg, Manitoba R3B ON2, Canada.

Journal of The Lepidopterists’ Society
33(2), 1979, 148-149

BISTON COGNATARIA (GEOMETRIDAE): FREQUENCY OF MELANIC
MALES IN TYRINGHAM, MASSACHUSETTS, 1958-1977

Sargent (1974, J. Lepid. Soc. 28: 145-152) reported the frequency of the melanic
versus the typical form of the Salt and Pepper Geometer, Biston cognataria (Guenée),
in collections totalling 129 specimens (presumably all male) from central Massachu-
setts for the years 1971-1973. The percent of melanics ranged from zero in 1971
to 5.6 in 1973. The overall incidence for the three years was 6/129 = 4.4 percent.
Sargent discussed the question of industrial melanism and urged the continued re-
porting of data bearing on this problem.

During 18 years of the period from 1958 through 1977 (omitting 1961 and 1962)
I examined a total of 833 males of this species taken at light in Tyringham, a rural
area in southern Berkshire County of western Massachusetts, elevation 313 meters.
Atmospheric pollution in the area was minimal as evidenced by the common presence
of lichens on tree trunks. Collections in each year covered a period from late May
to mid September or later. The overall incidence of melanism was 83/833 = 9.96
percent. The yearly incidence ranged from zero in six of the 18 years, to a high of
33.3 percent (2/6) in 1971 (Table 1). The average of yearly incidences was 7.31
percent. Abundance of moths available for examination ranged from zero in 1970
to 253 in 1966, in which year the incidence of melanics was 11.1 percent. If any
general trend is evident, it is toward a recent diminution in the number of melanics.

VoLUME 33, NUMBER 2 149

TaBLE 1. Incidence of melanism in males of Biston cognataria in Tyringham,
Massachusetts, 1958-1977.

Year Melanic Form Typical Form Total Examined Percent Melanic

1958 1 28 29 oD

1959 12 93 105 11.4
1960 6 OM 43 14.0

1961 (no records)
1962 (no records)
1963 ik 8 9 L1.1

1964 2 23 25 8.0

1965 a 69 16 9.2

1966 28 225 253 11.1
1967 V7 IVA 188 9.4

1968 5 ol 36 Nes)

1969 0 3 3 0.0
1970 0 0 0) —-

ial 2, 4 6 33.3

1972 0 4 4 0.0

1973 0 Tf 7 0.0

1974 0 10 10 0.0

1975 0 10 10 0.0
1976 2 10 12 NGC

1977 0 Wy li 0.0
Total 83 750 833 TB

AsHER E. TREAT, Research Associate, The American Museum of Natural History,
New York, New York 10024.

Journal of The Lepidopterists’ Society
33(2), 1979, 149-150

NOTES ON THE OCCURRENCE OF ERORA LAETA (LYCAENIDAE) IN
MICHIGAN’S WESTERN UPPER PENINSULA

Erora laeta (Edwards) has been known from only one location in the upper Great
Lakes region. On 14 May 1955, Edward G. Voss and Warren H. Wagner, Jr.
had the good fortune of encountering four individuals of this elusive butterfly, two
males and one female of which were captured. These records from Bliss Township,
Emmet Co., Michigan represented a first for the state and substantially extended
the known range of this exceedingly rare species. In a paper recounting these
captures (Voss & Wagner 1956, Lepid. News 10: 18-24), the authors gave consider-
able attention to the environmental characteristics of the collecting site. In brief,
all of the specimens were captured at or near a moist spot, which was fed by a small
ground spring situated on the south side of an east-west dirt section road. Botanical-
ly, the surrounding area was described as a very young deciduous woods, with
American beech (Fagus grandifolia Ehrh.), sugar maple (Acer saccharum Marsh. ),
basswood (Tilia americana L.), American elm (Ulmus americana L.), white ash
(Fraxinus americana L.), and pin cherry (Prunus pensylvanica L.) dominating.

150 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Beaked hazelnut (Corylus cornuta Marsh.) was not noted anywhere near and conifers
were conspicuously absent. Subsequent collecting at this site by Voss and Wagner,
along with numerous others, has failed to turn up additional specimens.

On 27 May 1975, I was collecting in an area SE of Porcupine Mountains State
Park, in Ontonagon Co., principally in search of colonies of Pieris virginiensis. Rich
northern hardwood forests along a spur service road to the Bergland Lookout Tower
provided excellent collecting for several spring desirables. At approximately 1730
(CDT), a small lycaenid was noted at a mudpuddle margin along this service road,
ca. 0.8 km ENE of and some 84 m below Bergland Tower (elevation 541 m).
Being immediately recognized as E. laeta, I wasted no time in netting it. The cap-
tured specimen was an immaculate female. After a brief period of disbelief and
celebration, other wet spots along the entire length of the service road were care-
fully checked five or six times, but no additional specimens were encountered. The
entire area was again scoured on the following day, but overcast weather conditions
greatly inhibited butterfly activity. Having been advised that weather conditions
were not expected to change for the better, I left the area, having lost hope of
finding additional individuals.

This site is approximately 400 km WNW of the Emmet Co. location where it
was found in 1955, thus representing the most northwestern station for E. laeta
known. Botanically the Ontonagon Co. site is quite dissimilar from that in Emmet
Co. It can be typified as a well drained, secondary northern hardwood forest,
dominated by basswood (Tilia americana L.), sugar and mountain maples (A. sac-
charum and spicatum Lam., respectively), red oak (Quercus rubra L.), and American
elm, with occasional balsam fir (Abies balsamea L.), intermixed with wetter areas
containing black ash (Fraxinus nigra Marsh.), and with trembling aspen (Populus
tremuloides Michx.) colonies throughout. Understory shrubs include beaked hazel-
nut and thimbleberry (Rubus parviflorus Nutt.). The most conspicuous herbs during
late May were the large-leaved aster (Aster macrophyllus L.), winter cress (Bar-
barea vulgaris R. Br.), nodding trillium (Trillium cernuum L.), sensitive fern
(Onoclea sensibilis L.), and several species of violets. Many additional weedy plants
occur along the disturbed roadside.

Other butterflies found associated with E. laeta at the Ontonagon Co. station in-
cluded Erynnis icelus (scarce; fresh), Amblyscirtes samoset (common; freshly
emerging), Carterocephalus palaemon (scarce; emerging), Papilio glaucus canadensis
(common; fresh), Pieris virginiensis (common; mostly worn), Celastrina argiolus
pseudargiolus (abundant; worn), Nymphalis vau-album (scarce; fresh), Polygonia
comma (common; fresh), and P. faunus (common; fresh). Judging from this isolated
capture, the peak flight period of laeta in the area appears to be fixed between the
full flights of the principally early spring species (P. virginiensis and C. argiolus
pseudargiolus) and those of late spring (A. samoset and C. palaemon).

Both American beech and beaked hazelnut are suspected larval hostplants of E.
laeta. American beech is abundant at the Emmet Co. collecting site, while beaked
hazelnut is very rare or altogether absent. At the Ontonagon Co. site, the opposite
is true: beaked hazelnut constitutes a dominating element, while American beech
was not noted at all. Warren H. Wagner, Jr., informs me (in litt.) that the western-
most extent of American beech in North America is in the north in eastern Marquette
Co., eastern Iron Co., and then extending down the eastern one-fifth of Wisconsin.
The nearest localities for beech are thus roughly 160 km east of the Ontonagon
E. laeta site. Most Midwest collectors have in the past associated E. laeta with
northern hardwood forests dominated by American beech but, in view of the present
capture, it would be advisable to also concentrate collecting efforts in northern hard-
wood situations as characterized above.

I wish to thank Dr. Warren H. Wagner, Jr., of the University of Michigan for
providing plant determinations and for reviewing the manuscript.

DanieL P. Oostinc, 7529 Walnut Avenue, Jenison, Michigan 49428.

VOLUME 33, NUMBER 2 Iisyll

Journal of The Lepidopterists’ Society
33(2), 1979, 151-152

EURISTRYMON ONTARIO (LYCAENIDAE): FIRST REPORT IN MICHIGAN

On 28 June 1975, Harvey and Oosting were collecting in an area NNW of
Morenci, Lenawee Co., Michigan, principally in search of Satyriwm caryaevorus. A
late afternoon stop was made at a likely-looking hairstreak location just south of Lime
Creek along Munson Highway, a secondary road running north-south along the sec-
tion line. An overcast sky had all but put an end to the day’s field work, but a quick
check was made of a roadside patch of mixed white and yellow sweet clover
(Melilotus alba Desr. and M. officinalis (L.) Lam., respectively ).

What appeared to be another Satyrium was noticed by both Harvey and Oosting
taking nectar from white sweet clover. It seemed to be somewhat different from
other hairstreaks taken during the day, and it was routinely swept up in Oosting’s
net for examination. Imagine our surprise to discover that it was a slightly worn
male ontario, the first collection for the state of Michigan. An immediate vigorous
search of the area yielded no additional specimens. The following day, accompanied
by Wagner, the collectors again investigated the entire area. Sunny skies provided
excellent Satyrium collecting, but additional specimens of E. ontario were not seen.
Nevertheless we made a careful survey of the habitat in terms of plant species, in
the hope that it might contribute something to our understanding of this elusive
butterfly and its occurrence in north-central and northeastern North America.

The roadbank patch of white and yellow sweet clover, where ontario was
captured, is bordered on the south by a large weedlot, including mainly Eurasian
plants which, like the sweet clovers, have become naturalized to a greater or lesser
extent as weeds. Southeast of the spot where ontario was collected is an extensive
cornfield, like many others in an area which is made up of farms interspersed with
mainly small wooded areas, these located primarily along stream valleys. Just to the
south of the locality is Lime Creek, running east-west. The stream valley is rather
heavily wooded, especially to the west of the locality (across the road), where there
is an old pasture woods with an extensive development of hawthorns (Crataegus
spp.) on the edges and in openings. The pastured forest (now including only a couple
of cattle, but formerly more) has a tall, heavy canopy made of very common plants
of sugar maple (Acer saccharum Marsh. ), black maple (A. nigrum Michx.), shagbark
hickory (Carya ovata (Mill.) K. Koch), white ash (Fraxinus americana L.) and
basswood (Tilia americana L.). Less common species include red maple (A. rubrum
L.), false shagbark (C. ovalis Wang.), American beech (Fagus grandifolia Ehrh.),
hop hornbeam (Ostrya virginiana Mill.) and red oak (Quercus rubra L.). In the
understory, the clonal shrub, prickly ash (Xanthoxylum americanum Mill.), is com-
mon and in the low herbaceous growth are recognized such rich forest species as
Virginia snakeroot (Aristolochia serpentaria L.), may apple (Podophyllum peltatum
L.) and bloodroot (Sanguinaria canadensis L.). Lopseed (Phryma leptostachya L.)
is especially common as an herbaceous forest-floor plant.

There is a step, darkly shaded slope on the north, running down to the stream bed
and an alluvial forest, where such woody species as pawpaw (Asimina_ triloba
(L.) Dunal), spicebush (Lindera benzoin L.), sycamore (Platanus occidentalis L.),
and cottonwood (Populus deltoides Marsh.) are frequent to common. On the flood-
plain forest floor are found such rich-soil plants as creeping fragile fern (Cystopteris
protrusa (Weath.) Blasdell), blue cohosh (Caulophyllum thalictroides L.), liverleaf
(Hepatica acutiloba DC) and wood nettle (Laportea canadensis L.). Persons
familiar with southern Michigan vegetation will recognize here a plant association
of luxuriant, rich woods—hardly comparable to a “shale barrens” like that described
by Clench (1971, J. Lepid. Soc. 25: 80-82).

152 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

It is possible that the specimen of ontario was blown in or flew in from some
distance away. Judging from its condition, however, the specimen might have
originated near by. Indeed the presence of the suggested larval foodplant genus,
Quercus, makes this not unlikely. Other species of hairstreaks, particularly S. calanus
falacer, were common in light gaps in the woods, where they perched on shrubs and
small trees. Presumably the falacer are attracted into open, weedy areas by the
flowers present there, and they return to their natural woodland and woods-edge habi-
tats when they are not feeding. In addition to S. c. falacer, the following species of
butterflies were found in the general area: Wallengrenia otho egeremet (common),
Polites coras (scarce), Papilio glaucus (common), P. troilus (scarce), Satyrium
caryaevorus (common), S. acadica (scarce), Chlosyne nycteis (scarce) and Speyeria
cybele (common).

DANIEL P. Oostinc, 7529 Walnut Avenue, Jenison, Michigan 49428.

Donatp J. Harvey, Department of Zoology, University of Texas, Austin, Texas
(0012.

WarreN H. Wacner, Jr., Department of Botany, University of Michigan, Ann
Arbor, Michigan 48109.

Date of Issue (Vol. 33, No. 2): 20 July 1979

EDITORIAL STAFF OF THE JOURNAL
AUSTIN P. PLATT, Editor ;
Department of Biological Sciences
University of Maryland Baltimore County, 5401 Wilkens Avenue
Catonsville, Maryland 21228 U.S.A.
FRANCES S. CHEW, Managing Editor

Department of Biology
Tufts University
Medford, Massachusetts 02155 USA

| Douc.as C. FERGUSON, Associate Editor THEODORE D. SARGENT, Associate Editor

NOTICE TO CONTRIBUTORS
Contributions to the Journal may deal with any aspect of the collection and study of

, _ Lepidoptera. Contributors should prepare manuscripts according tothe following in-

structions.
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Literature Cited: References in the text of articles should be given as, Sheppard
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_ the heading LITERATURE CITED, in the following format:

fe SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd. ed. Hutchinson, London.
5 209 p.

196la. Some contributions to population genetics resulting from the study of
the Lepidoptera. Adv. Genet. 10: 165-216.

In the case of general notes, references should be given in the text as, Sheppard (1961,
Adv. Genet. 10: 165-216) of (Sheppard 1961, Sym. R. Entomol. Soc. London 1: 23- -
30).

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CONTENTS

SUBSPECIFIC VARIATION IN BUTTERFLIES: ADAPTATION AND
DISSECTED POLYMORPHISM IN PIERIS (ARTOGEIA) ( PIERIDAE).
S. R: Bowden

NOTES ON THE LIFE CYCLE AND NATURAL HISTORY OF
BUTTERFLIES OF EL SALVADOR. III. C. HisToriIs opIUs AND
COEA ACHERONTA (NYMPHALIDAE—COLOBURINAE). Albert
Muyshondt, Jr. & Alberto Muyshondt ____--_-.-.-----------_--__—

REVIEW OF THE MEXICAN POLYTHRIX WATSON 1893 (HESPERI-

IDAE). H. A. Freeman 0

PREDATORY BEHAVIOR IN LITHOPHANE QUERQUERA AND OTHER
SPRING CATERPILLARS. Dale F. Schweitzer ________.._..._____.

A HOLOTYPE DESIGNATION FOR PAPILIO CINYRAS RIDENS MASTERS
1971 (PAPILIONIDAE). John H. Masters _.____. > ae

THE TYPE LOCALITY OF ARGYNNIS ZERENE BOISDUVAL (NYM-
PHALIDAE): A CORRECTION. John H. Masters ____-_....__-

NOTES ON CHILEAN OECOPHORIDAE. J. F. Gates Clarke 333

GENERAL NOTES

Melipotis indomita (Noctuidae) in Hawaii and Connecticut. Dale F.
Schweitzer 2.220220

Notes of Maryland Lepidoptera. 6. Occurrence of Boloria selene (Nym-_
phalidae) in Maryland. William A. Andersen ¢& Robert S. Simmons --
Two southern cossids (Cossidae) in the New Jersey pine barrens. Dale F.
Schweitzer 2.000

A melanistic specimen of Antheraea polyphemus polyphemus (Saturni-
idae). William B. Preston & Wm. Brian McKillop __-___----------------

Biston cognataria (Geometridae): frequency of melanic males in Tyring-
ham, Massachusetts, 1958-1977. Asher E. Treat _____--__.----_------_-:

Notes on the occurrence of Erora laeta (Lycaenidae) in Michigan’s western
upper peninsula. Daniel P: Oosting _..._..0...... eee

Euristrymon ontario (Lycaenidae): first report in Michigan. Daniel P.
Oosting, Donald J. Harvey, & Warren H. Wagner, Jr. _-----------------

BOOK REVIEW 22000000 as

Volume 33 1979 © Number 3

JOURNAL

of the

LEPIDOPTERISTS’ SOCIETY

Published quarterly by THE LEPIDOPTERISTS’ SOCIETY

Publié par LA SOCIETE DES LEPIDOPTERISTES
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN
Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS

29 October 1979

THE LEPIDOPTERISTS’ SOCIETY

EXECUTIVE COUNCIL

T. D. SARGENT, President I. F. B. COMMON, Immediate Past
A. M. SHAPIRO, Ist Vice President President
ATUHIRO SIBATANI, Vice President JULIAN P. DONAHUE, Secretary

RONALD LEUSCHNER, Treasurer

Members at large:

J. F. EMMEL C, D. FERRIS M. DEANE BOWERS
R. R. GATRELLE J. Y. MILLER E. HODGES
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mally constituted in December, 1950, is ““to promote the science of lepidopterology in
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facilitate the exchange of specimens and ideas by both the professional worker and the
amateur in the field; to secure cooperation in all measures” directed towards these
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Cover illustration: Third instar larva of Limenitis archippus Cramer (Nymphalidae)
preparing to enter winter diapause. The larva is resting on the lip of its hibernaculum
constructed from the basal portion of a chewed tubular willow leaf (Salix babylonica
Linnaeus) covered with silk. In the autumn such larvae begin facultative diapause in
response to decreasing day-length. Original drawing by Mr. George C. Ford, Jr., Graph-
ics Illustrator, Department of Biological Sciences, University of Maryland Baltimore
County, 5401 Wilkens Avenue, Catonsville, Maryland 21228.

pecicaire Pea

reat

><

°° 2 ae Decal ee ee ee

Py ke oe ee Sie i eg

JOURNAL OF

Tue Leprporpreristrs’ SOCIETY
Volume 33 1979 Number 3

Journal of the Lepidopterists’ Society
33(3), 1979, 153-161

THE MALAISE TRAP AS A MEANS OF SAMPLING
BUTTERFLY POPULATIONS IN KENTUCKY!

CHARLES V. COVELL, JR.
Dept. of Biology, University of Louisville, Louisville, Kentucky 40208

AND

PAUL H. FREYTAG
Dept. of Entomology, University of Kentucky, Lexington, Kentucky 40506

ABSTRACT. Butterflies representing 57 species in 9 families were collected by
means of Malaise traps at 10 localities in Kentucky during the summers of 1970-1977.
The species are listed along with collecting sites, months of capture, and numbers of
individuals caught. Discussion of the potential of Malaise traps for sampling butterfly
populations includes an analysis of two of the three broods of Phyciodes tharos (Drury)
at Lexington, Kentucky as sampled by Malaise traps in 1971.

One of the problems in carrying out qualitative and quantitative
studies of butterfly populations in a given area is capturing specimens.
Butterflies, often highly mobile and difficult to approach, have tradi-
tionally been stalked with net and killing jar—a very time-consuming
procedure. Some species—those that can be attracted to rotten fruit
and other non-nectar adult foods—can be collected in bait traps but
this technique can be used for only a small proportion of species in
most butterfly communities. The use of pheromone traps for butter-
flies is at best in its infancy, and would present similar limitations in
breadth of appeal as the bait trap. One promising technique is the
Malaise trap (Fig. 1), which collects insects by blocking their passage
and taking advantage of their instinct to climb upward. This trap has
already been modified by various workers for different purposes but

1 Contribution No. 192 (New Series) of the Department of Biology, University of Louisville. This paper (78-7-
170) is published with the approval of the Director of the Kentucky Agricultural Experiment Station, Lexington,
Kentucky 40506.

154 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

apparently this paper is among the first to report its use in sampling
butterfly populations.

Townes (1962, 1972) adapted the Malaise trap for the purpose of
collecting large numbers of Hymenoptera for taxonomic purposes.
Others have followed, either to assess total insect communities
(Geijskes, 1968; Matthews & Matthews, 1970), or to collect particular
insect groups (Breeland & Pickard, 1965, for mosquitoes; Adkins et
al., 1972, for Tabanidae). Lepidoptera are usually represented as a
percentage of the whole catch in general insect surveys. Percentages
of each order are based on a count of all members of that order in the
sample container. Thus moths and butterflies are generally lumped
together, and rarely are they separated even to family level. Geijskes
(1968) found in Surinam that Lepidoptera were 14.3% of catches in
1963-64, but seasonal fluctuations of 8-37% were noted. Matthews &
Matthews (1970) found that Lepidoptera constituted 7.2% of their to-
tal catch during 13 weeks in 1967 at Rensselaerville, New York. Their
Lepidoptera were “almost exclusively moths, the bodies of over half
(49%-69%) measuring less than 3.0 mm excluding antennae.”

One comprehensive study of butterfly populations collected by
Malaise trap was that of Owen (1971), who used this technique in
Kampala, Uganda, in 1965, and at Freetown, Sierra Leone, in 1968.
He showed monthly fluctuations in numbers of individuals collected,
grouping his catch by family. His only tabulation at the species level
was Acraea bonasia (Fabr.). That common butterfly was present all
year, but showed a definite peak in February and March (Owen, 1971:
70). While Malaise trap samples played only a small part in his overall
study, Owen was possibly the first to demonstrate the value of Malaise
traps in sampling butterflies. More recently Walker (1978) has dem-
onstrated its value as a technique for studying butterfly migration.

METHODS

In 1970 the junior author, a leafhopper specialist, began a program
of Malaise trap sampling at several localities in Kentucky: Lexington
(Fayette Co.); Glendale (Hardin Co.); Bardstown (Nelson Co.); Rob-
inson Forest (Breathitt Co.); Princeton (Caldwell Co.); Paris (Bourbon
Co.); Spears (Jessamine Co.); and rural areas in Pulaski and Wayne
counties. The junior author separated the Lepidoptera from other or-
ders and the senior author subsequently identified them for the Ken-
tucky Lepidoptera Survey. Although the Lepidoptera collected were
mainly moths, a large number of small and medium-sized butterflies
appeared in the samples. This material was augmented by specimens
collected by University of Louisville graduate students under the su-
pervision of the senior author in Bernheim Forest (Bullitt Co.).

VOLUME 33, NUMBER 3 155

Fic. 1. Malaise trap at Bernheim Forest, Bullitt County, Kentucky, 1977. The trap
was 6 ft long, 4 ft wide, and 6 ft 8 in high at one end. At the high point, a hole of 2%
in diameter led into a plastic jar, to which a plastic cyanide jar could be screwed by
means of lids glued back to back and drilled with a similarly large opening to let the
insects through.

Two variations of the Malaise trap were employed to obtain the
results reported here. The traps used by the junior author at all sites
except Bernheim Forest were Model 300.5 from Survival Security
Corporation, Lake City, Minnesota 55041 (not illustrated). About 7 ft
high, these traps consisted of dark nylon netting in four panels rising
to a central peak supported by an aluminum pole. The top of the trap
had an inverted plastic cone to direct the insects into a smaller cone
placed snugly beneath the first, then narrowing downward into a cy-
anide killing jar. These components were supported by the center
pole. Entrance space between the two cones was provided by cuts
about % inch deep around the top of the inner cone. The traps were
selective for small insects, especially leafhoppers, so few large but-
terflies were included in the samples.

The trap used at Bernheim Forest (Fig. 1) was built by University
of Louisville graduate student Larry Canterbury according to the di-
rections of Townes (1972). The openings into the killing jar were
larger in this trap, and several of the large butterfly species listed in
Table 2 were taken only at the Bernheim Forest site. The Bernheim
trap was encircled by barbed wire to discourage tampering.

156 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TABLE 1. Numbers of total butterfly species of each family collected by Malaise trap
in Kentucky, 1970-1977. Percentages of the total are indicated in the right-hand col-
umn.

Number of species

Family Kentucky total Malaise traps Percentage
Hesperiidae 44 21 48
Papilionidae 8 3 Ona
Pieridae 13 6 46.2
Riodinidae 2 0) 0
Lycaenidae NS) 9 36
Libytheidae 1 ] 100
Nymphalidae 29 13 45
Danaidae Il 1 100
Satyridae 8 3 Oo
Total butterfly species 131 57 43.5

RESULTS

The butterflies collected in Malaise traps were identified by the
senior author and named according to the current checklist for North
American butterflies (Dos Passos, 1964) and later modifications such
as Howe (1975). Covell (1974) listed 123 butterfly species from Ken-
tucky, and since then 8 more have been added. As of this writing, 57
of the 131 species, or 43.5%, have been collected in Malaise traps. No
additions to the state list have resulted yet from these samples, but
substantial numbers of new county records and some additional sea-
sonal information has been recorded.

Table 1 indicates the Malaise-trapped butterfly species in each fam-
ily compared with the total number known from Kentucky in that
family. Species that have not been collected include (1) 5 species that
are clearly “strays” and known from single Kentucky records; (2) 23
species that are either very rare, or which are found in very restricted,
specialized habitats not yet sampled by Malaise trap; or (3) some
of the very large butterflies which were not caught despite the Bern-
heim trap with large opening. It may also be that flight and walking
habits of some species, not yet appreciated, may account for their
absence from samples. Only one family occurring in Kentucky, the
Riodinidae, has not been taken by Malaise trap; and these fly for short
periods in very specialized habitats.

Table 2 lists all the butterfly species collected by Malaise trap in
Kentucky from 1970 through 1977. Although the number of specimens
collected by this technique over the eight-year period is only 1,883,
one must remember that except at Bernheim Forest the traps were
not designed or placed for butterfly collecting. The majority of spec-

VOLUME 33, NUMBER 3 157

TABLE 2. List of species of butterflies in each family collected by Malaise trap in
Kentucky, 1970-1977. To the right of each species name are numbers representing
localities from which each was collected (1-10), the months of capture (1-12), and the
total number of specimens taken. Localities by number are as follows: 1, Bernheim
Forest, Bullitt Co.; 2, Lexington, Fayette Co.; 3, Glendale, Hardin Co.; 4, Spears,
Jessamine Co.; 5, rural Mercer Co.; 6, Robinson Forest, Breathitt Co.; 7, Princeton,
Caldwell Co.; 8, Bardstown, Nelson Co.; 9, rural Wayne Co.; and 10, Paris, Bourbon
Co.

Speci-
Species Localities Months mens

HESPERIIDAE
Amblyscirtes hegon (Scudder) 1 5 2D
Euphyes vestris metacomet (Harris) 2 5-9 42
Poanes zabulon (Bdv. & LeConte) 1,2,6,10 5-9 8
Poanes hobomok (Harris) 1 5=6 3
Atrytone delaware (Edwards) 2 6-9 8
Atalopedes campestris (Bdv.) 2,4,5,8,10 7-10 87
Pompeius verna (Edw.) 2 6,8-9 9
Wallengrenia egeremet (Scudder) 2,4,8,10 6-9 4]
Polites coras (Cramer) 1-4.8 5-10 102
Polites themistocles (Latreille) 1-6,.8 5-9 169
Polites origenes (Fabr.) 1,2,6,8 6,8 8
Thymelicus lineola (Ochs.) 2,4 5,6 68
Ancyloxypha numitor (Fabr.) 2-5,10 6-9 13
Nastra lherminier (Latreille) 45,7 7-8 6
Pholisora catullus (Fabr.) 2-5.8 5-9 214
Pyrgus communis (Grote) 2.3.8 7-10 15)
Erynnis icelus (Scudder & Burg.) 2:5 4-8 50
Erynnis brizo (Bdv. & LeConte) 1 3-4 31
Erynnis horatius (Scudder & Burg.) 6 7 1
Erynnis juvenalis (Fabr.) 1 3-4 26
Epargyreus clarus (Cramer) 1,2,5,6,8 46-8 18

PAPILIONIDAE
Battus philenor (Linn.) 1 4 1
Papilio troilus Linn. 1 4.6 5
Graphium marcellus (Cramer) 1 4.6 3

PIERIDAE

Pieris rapae (Linn.) 2-5,8 5-9 139
Colias eurytheme Bdv. 1-5,10 5-8, 10 29
Colias philodice Godart 1-5 46-10 36
Eurema lisa Bdv. & LeConte 3 a 1
Eurema nicippe (Cramer) 335,21 7-8 2
Anthocharis midea Hibn. 1 3-4 4

LYCAENIDAE
Harkenclenus titus mopsus (Hubn.) 2 6 1
Satyrium calanus falacer (Godart) 4,5 6,7 2
Calycopis cecrops (Fabr.) 6 5 1
Callophrys henrici (Grote & Rob.) 1 3-4 )
Strymon melinus Hubn. 1-3,5,8 4-9 21
Lycaena hyllas (Cramer) 3 7 1
Lycaena phlaeas americana Harris 2-4. 6-9 19
Everes comyntas (Godart) 1-5,8 4-9 84

A ee ee an

158 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

TABLE 2. Continued.

Speci-
Species Localities Months mens

Celastrina argiolus pseud-

argiolus (Bdv. & LeConte) 1 4 6

LIBYTHEIDAE
Libythaena bachmanii (Kirtland) Drake 6-8 8
NYMPHALIDAE
Asterocampa celtis (Bdv. & LeConte) 1-4,7,10 6-9 58
Asterocampa clyton (Bdv. & LeConte) 2,3,7,8,10 8-9 8
Limenitis arthemis astyanax (Fabr.) 3 qf 1
Vanessa atalanta (Linn.) 2 5,6 5
Vanessa virginiensis (Drury) 2 5,6 1
Precis coenia (Hubn.) 3,8,9 6-9 4
Polygonia interrogationis (Fabr.) 1,2 5,6,8 6
Polygonia comma (Harris) 1 4 il
Chlosyne nycteis (Doubleday) 1,3,4 5-9 10
Phyciodes tharos (Drury) 1-3,5,6,8 4-10 379
Boloria bellona (Fabr.) 2 6-9 69
Speyeria cybele (Fabr.) eZ 6-7 7
Euptoieta claudia (Cramer) 263 6-9 ¢
DANAIDAE
Danaus plexippus (Linn.) 3 8 1
SATYRIDAE
Cyllopsis gemma (Hubn.) 6 6-9 4
Euptychia hermes sosybius (Fabr.) 6 6-9 IL)
Megisto cymela (Cramer) 1,6 5-7 18
Totallesss

imens were small species, the common Phyciodes tharos (Drury) con-
stituting 20% of the total. Its wingspan is approximately 3 cm. Also,
the traps were placed in open fields and meadows with two excep-
tions: the Robinson Forest and Bernheim Forest sites. The few Sa-
tyridae collected can be attributed largely to that factor, since most of
the Kentucky members of this family are woodland species. Further-
more, although some early spring collecting was done, the period
from mid-March to the end of April was largely neglected; and several
early spring butterflies were not collected—especially those restricted
to woodland habitats. While only months of capture are indicated
here, further study of the data would reveal trends in abundance of
given species at a particular locality during a given year. These data
might be valuable in understanding population dynamics, especially
when correlated with climatic factors such as temperature and rainfall.
An example of a more precise analysis is given in Fig. 2 for Phyciodes
tharos, which has three broods in Kentucky. At Lexington in 1971, 88

VOLUME 33, NUMBER 3 159

no FF Oo

APR. MAY Junt JULY J
AUG. SEP. O
23=30) ik =i 127i Pil=si0) Gl =6) Gil Dilezyo i =i II=20 20=HiL 1 =] 9=20 20=39 i =13 1B=Ail 21-30 ft

Fic. 2. Temporal distribution of Phyciodes tharos (Drury) collected in a Malaise
trap in a Lexington, Kentucky, meadow, 23 May-11 Oct. 1971.

individuals were collected during the 17 time increments indicated
on the graph. From prior experience we expect the three broods to
occur in spring, midsummer, and late summer to fall. We are not
certain if the single capture of 12-21 May represents the spring or
midsummer brood. The spring brood was not properly sampled since
that trap was not set up until about 20 April. The midsummer peak
at that site in 1971 was apparently during 1-11 July, with 24 individ-
uals collected. The next peak during the last 10 days of August and
the first 13 of September was also substantial. If the trap had been
checked daily instead of at rather irregular intervals, a more sensitive
picture of the dynamics of P. tharos could have been obtained. Sim-
ilar data from several years would be interesting, to give comparisons
of both abundance and peak flight times for adult P. tharos. The Mal-
aise technique can be used to remove individuals permanently from
the population by use of a killing jar; or individuals can be marked,
released, and recaptured on succeeding days for studies of that nature.

160 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Since Fig. 2 was constructed in hindsight from butterflies collected
by the junior author in 1971, one should not assume that it is intended
to give anything more than a rough picture of P. tharos dynamics at
the Lexington site that summer.

CONCLUSIONS

Large numbers of insects, including such fast fliers as moths and
butterflies, can be collected with relatively little effort by using Mal-
aise traps. By experimentation with the basic design of the trap one
can enhance the efficiency of capture of either butterflies in general,
or certain desired species. Some of the variables which still need
attention include netting color, texture, and durability; trap size and
shape; placement of collecting container, size and shape of opening
into container; dimensions of container itself. Position of the trap in
suitable habitat is also essential to maximize collection. Placement in
a butterfly fly-way through a forest is an example of proper choice in
this respect. Addition of bait may increase captures of certain desired
species. If killed specimens in excellent condition are required, one
must seek a long-lasting killing agent of strength sufficient to prevent
damage by beetles and large insects (including Lepidoptera) that get
into the container. A large live-catch cage could also be used, espe-
cially if one does not wish to make a general survey or kill everything
that enters the trap. By using such a cage one can free insects that
one is not interested in keeping, and thus avoid needless killing of
large numbers of other insects. The more frequently the trap is emp-
tied, the better one can expect specimen condition to be. In the col-
lections studied in this survey, however, the condition of butterflies
was remarkably good; a large proportion was nearly perfect and good
enough to put in a collection. A major drawback to using killed spec-
imens from Malaise traps seems to be the unpreventable problem of
wings folding down beneath the body as the insect dies. Such spec-
imens often have drooping wings, even after a good job of relaxing
and spreading. Experimentation with other killing agents may alle-
viate this problem.

In addition, some other negative aspects of Malaise trap collecting
in general include:

(1) Expense: Unless one makes the trap himself, commercially avail-
able models run from $100 to more than $300.

(2) Weather: Rain may ruin samples; wind can knock down the trap
or bring tree limbs down on it; and sun, rain, and wind can cause
deterioration of some netting materials, e.g., nylon. Dacron is
more durable according to Townes (1972).

VOLUME 33, NUMBER 3 161

(3) Vandalism: Despite signs and even barbed wire fencing (Fig. 1),
traps are often prey to vandals in isolated localities. We know of
cases in which traps have been destroyed or stolen on seemingly
protected property.

(4) Lack of selectivity: As mentioned above, one is likely to kill large
numbers of insects indiscriminately by this technique unless a
live-catch container is used and the trap is emptied daily.

Despite these shortcomings, however, Malaise traps show great prom-
ise for a wide variety of insect population studies, and are increasingly
used throughout the world. This survey has added much information
to the current study on butterflies of Kentucky; the moth material,
when identified, should provide numerous important records for that
aspect of the survey.

ACKNOWLEDGMENTS

The authors wish to thank their student assistants and colleagues
for help in operating the Malaise traps in the various localities, and
for sorting the samples: D. P. Beiter, A. J. Brownell, L. E. Canterbury,
W. B. Early III, G. F. Florence, C. H. Kaster, J. S. Lesshafft, Jr., J.
Pohlman, H. G. Raney, S. Reigler, and C. K. Sperka. We are also
grateful to the Isaac Bernheim Foundation, Louisville, Kentucky, for
the funding to carry out the survey of Bernheim Forest.

LITERATURE CITED

ADKINS, T. F. Jr., W. B. EZELL JR., D. C. SHEPPARD & M. M. ASHLEY JR. 1972. A
modified canopy trap for collecting Tabanidae (Diptera). J. Med. Entomol. 9(2):
183-185.

BREELAND, S. G. & E. PICKARD. 1965. The Malaise trap—an efficient and unbiased
mosquito collecting device. Mosquito News 25: 19-21.

COVELL, C. V. JR. 1974. A preliminary checklist of the butterflies of Kentucky. J.
Lepid. Soc. 28(3): 253-256.

Dos Passos, C. F. 1964. A synonymic list of the Nearctic Rhopalocera. Mem. Lepid.
Soc. 1: 1-145.

GEIJSKES, D. C. 1968. Insect collecting in Surinam with the help of “Malaise” traps.
In Studies on the fauna of Surinam and other Guyanas, no. 39. Natuurwetensch.
Studiekring Surinam Ned. Antillen 48: 101-109.

Howe, W. H. 1975. The butterflies of North America. Doubleday, Garden City, N.J.
63a)ps097 pl.

MATTHEWS, R. W. & J. R. MATTHEWS. 1970. Malaise trap studies of flying insects in
a New York mesic forest. I. Ordinal composition and seasonal abundance. J. New
York Entomol. Soc. 78: 52-59.

OWEN, D. F. 1971. Tropical butterflies. Clarendon Press, Oxford. 214 pp.

TOWNES, H. 1962. Design fora Malaise trap. Proc. Entomol. Soc. Washington 64: 253-
262.

1972. A light-weight Malaise trap. Entomol. News 83: 239-247.

WALKER, T. J. 1978. Migration and re-migration of butterflies through northern pen-
insular Florida: Quantification with Malaise traps. J. Lepid. Soc. 32(3): 178-190.

Journal of the Lepidopterists’ Society
33(3), 1979, 162-166

DIFFERENTIAL GROWTH AMONG LARVAE OF CITHERONIA
REGALIS (SATURNIIDAE) ON THREE
GENERA OF FOODPLANTS

C. BROOKE WORTH
R. F. D. Delmont, New Jersey 08314

THOMAS F. WILLIAMS, AUSTIN P. PLATT, AND BRIAN P. BRADLEY

Department of Biological Sciences, University of Maryland Baltimore County,
5401 Wilkens Ave., Catonsville, Maryland 21228

ABSTRACT. Larvae of Citheronia regalis reared under similar outdoor conditions
grow both larger and more rapidly on persimmon than on either sweetgum or sumac.
We suggest that the relationship of C. regalis to persimmon is one of long standing,
originally established in the Neotropics, whereas both sweetgum and sumac may have
been exploited secondarily, as the moth entered temperate regions in more recent
times.

Many species of Lepidoptera are polyphagous in the larval stage.
For these larvae there often exists a spectrum of foodplant suitability,
including “preferred” and “acceptable” types of host plants. Such
preferences have a biochemical basis through which larvae respond
to chemosensory cues. During the course of evolution, the longer an
herbivorous insect and a particular plant species have been associ-
ated, the more compatible the two are likely to have become, both
biochemically and nutritionally, as each species influences the evo-
lution of the other (Brower & Brower, 1964; Ehrlich & Raven, 1965).

Larvae of Citheronia regalis Fabricius feed on a wide variety of
deciduous trees and shrubs, including as preferred foodplants: black
walnut (Juglans nigra Linnaeus), butternut (J. cinerea L.), hickories
and pecan (Carya spp.), persimmon (Diospyros virginiana L.), sweet-
gum (Liquidambar styraciflua L.), as well as mountain, smooth, stag-
horn, and wing-rib sumac (Rhus spp.). Alternate foodplants include
ashes (Fraxinus spp.), blackgum (Nyssa sylvatica Marsh), oak (Quer-
cus spp.), sycamore (Platanus occidentalis L.), willow (Salix spp.),
lilac (Syringa vulgaris), and even cotton and sea-island cotton (Gos-
sypium spp.), according to Holland (1903), Villiard (1969), and Fer-
guson (1971). This paper reports differential growth and development
times of C. regalis reared on persimmon, sweetgum, and wing-rib
sumac (Rhus capallina L.). Furthermore, we will suggest possible
reasons why C. regalis larvae exhibit differential growth rates on
these three foodplant genera.

MATERIALS AND METHODS

Rearing was done out-of-doors on a farm in Eldora (Cape May Co.),
New Jersey by C.B.W. during the summer of 1977. Equal numbers of

VOLUME 33, NUMBER 3 163

16 PERSIMMON SWEETGUM

PUPAL WEIGHT (gq)

DAYS TO PUPATION —=—

Fic. 1. Linear regression of pupal weight vs. larval development time for Cithe-
ronia regalis on three foodplants. Solid circles are data for males; open circles denote
females. The overall slope is —0.28 (P < 0.001). Slopes of individual lines (except
Sweetgum 2 2 and Sumac ¢ ¢) alse are significant (P < 0.001).

eggs from two females were placed in fine-mesh cloth nets on branch-
es of the three host plants. Locations facing the same direction and
having the same temporal exposure to sunlight were chosen. Larvae
were reared communally until third instar, at which time they were

TABLE 1. Effects of foodplant and sex on pupal weights (g) of Citheronia regalis
larvae, with 2-way ANOVA Test and mean pair comparisons. (Sample sizes are given
in brackets.)

FOODPLANT
Persimmon (PR) Sweetgum (SG) Sumac (SM)
did O 4 30:38 [14 2 = OTT «6.3 0.645]
Pupal weight (x = S.E.) Oe te Oo Els) 2 95--=- 0602) 8-9== 0:53: [6]

ANOVA of pupal weights!
(food plant and sex are fixed):

Source of variation df Mean Square F-statistic P
Between foodplants 2, 45.2 15.9 <0.005
Between sexes 1 16.9 5.8 <0.01
Interaction Ye 4.8 1.6 >0.10 (NS)
Within subclasses 4] 2.9 — —

If the means (x) are ordered from largest to smallest, Duncan’s multiple range tests show that the following means
do not differ significantly from each other (P > 0.05): PRE 2, PRS d, SG22 do not differ significantly; PRd¢,
SG22,SM2°2,SG¢é¢ do not differ significantly; S66 $¢ and SMé¢ do not differ significantly.

164 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TABLE 2. Effects of foodplant and sex on development time (days) of Citheronia
regalis larvae with 2-way ANOVA test and mean pair comparisons. (Sample sizes are
given in brackets).

FOODPLANT
Persimmon (PR) Sweetgum (SG) Sumac (SM)

Development time 2° 38.3 + 0.83 [14] 42.4 + 1.25 [7] 48.2 + 1.93 [5]
(x == Sky) 33 39.8 + 0.96 [13] 43.5 + 1.50 [2] 50.3 + 1.28 [6]

ANOVA of development times
(foodplant and sex are fixed):

Source of variation df Mean Squares F-statistic P
Between foodplants 2 415.6 Som <0.005
Between sexes 1 27.6 2.4 =(.10 (NS)
Interaction y) 0.5 0.04 >0.25 (NS)
Within subclasses 4] IL 8} = —

Duncan’s Multiple range tests show that the means (sexes combined) differ significantly from each other:
PR < SG < SM (P < 0.05).

reared singly or in pairs, depending on their size. Further rearing was
done inside heavy cloth bags (pillow cases) covering terminal branch-
es. During the fifth instar all larvae were inspected daily to determine
the date each ceased feeding, and when each began to crawl around
inside its cloth bag. Larvae were then brought indoors and were al-
lowed to pupate in individual plastic boxes. The sex and pupal weight
of each were determined on the day after the larval—pupal ecdysis.

RESULTS

Although initially equal numbers of eggs were placed on the three
foodplants, the number of larvae which actually began to feed after
hatching on each plant type was not recorded. When the larvae were
transferred at third instar, there were 31 feeding on persimmon, 10
on sweetgum, and 13 on sumac. Thus, following the third instar, the
C. regalis larvae exhibited approximately equal survival rates through
pupation; these rates ranged from 85% on sumac to 90% on sweetgum,
with 87% survival on persimmon.

Our data have been analyzed using two-way analysis of variance,
with foodplant and sex fixed. In Table 1 the effects of foodplant
and sex on pupal weight both are significant, but the interaction
term is not. The Duncan’s Multiple Range Tests on data in Table
1 indicate which of the group means differ significantly from one
another. Table 2 shows that only foodplant affects development
time, with neither sex nor the interaction term being significant. Fig.
| presents a linear regression analysis of pupal weight vs. larval de-
velopment time on each of the three foodplants. An inverse relation-

VOLUME 33, NUMBER 3 165

ship between pupal weight and development time is shown by these
graphs. Clearly, the C. regalis larvae grew more rapidly on persim-
mon than on either sweetgum or sumac. Larval growth was retarded
4—5 days on sweetgum, and 10-11 days on sumac, as compared to
persimmon.

DISCUSSION AND CONCLUSIONS

These results suggest that persimmon may be a more favorable
foodplant for C. regalis in the northeastern United States than are
either sweetgum or sumac. Larvae fed on the two latter plants not
only exhibited a smaller final size, but they also required a longer
feeding period. Yet C. regalis is know to oviposit on all three host
plants in Eldora, New Jersey, and its larvae often have been found
feeding in the wild on them. The local population probably exploits
numerous other deciduous hostplants as well; both black walnut and
mockernut hickory (Carya tomentosa Nuttall) are common species on
the wooded areas nearby.

The geographic distribution of C. regalis closely matches the range
of persimmon, whereas its other foodplants (except for sweetgum)
have less extensive (mainly temperate) ranges (Fowells, 1965). We
may now ask what reasons account for persimmon’s optimal qualities
as a foodplant for C. regalis? Possibly, this represents a relationship
of long standing, whereas the contact between C. regalis and both
sweetgum and sumac may have occurred more recently.

Citheronia has its generic focus in Central and South America, and
persimmon, likewise, is a member of a predominantly tropical family,
the ebonies (Ebenaceae). All of the other preferred foodplants of C.
regalis are members of principally temperate zone families, either the
Juglandaceae (walnuts and hickories), the Hamamelidaceae (sweet-
gum), or the Ancardiaceae (sumacs). These foodplants also are shared
by Actias luna Linnaeus, another temperate member of predomi-
nantly Asiatic and tropical saturnid groups. Furthermore, the closest
taxonomic relatives of C. regalis common in temperate regions (C.
sepulcralis Grote and Robinson and Eacles imperialis Drury) are both
pine feeders and have exploited very different plant species.

Although our data are by no means sufficient to prove our hypoth-
esis we suspect that C. regalis and persimmon may have established
their relationship long ago in the tropics, and that the two only re-
cently invaded the North American temperate zone as biological con-
sorts. The spread of C. regalis to mainly temperate plants such as
sweetgum and sumac, then, may represent more recent relationships
which are still evolving. Further comparative foodplant studies of C.

166 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

regalis and its other host plant species are warranted to test this hy-
pothesis.

ACKNOWLEDGMENTS

We wish to thank Mr. T. Ford of the Department of Biological Sci-
ences, U.M.B.C. for technical assistance. We also are indebted to Dr.
R. S. Peigler of the Department of Entomology, Texas A&M Uni-
versity, for comments relating to these ideas.

LITERATURE CITED

BROWER, L. P. & J. V. Z. BROWER. 1964. Birds, butterflies, and plant poisons: A study
in ecological chemistry. Zoologica 49: 137-159 .

EHRLICH, P. R. & P. H. RAVEN. 1965. Butterflies and plants: a study in evolution.
Evolution 18: 586-608.

FERGUSON, D. C. 1971. In Dominick, R. B., et al., The moths of America north of
Mexico, Fasc. 20.2A, Bombycoidea (Saturniidae), E. W. Classey, Ltd., Middlesex,
England. 153 pp.

FOWELLS, H. A. 1965. Silvics of forest trees of the United States, Agriculture Hand-
book No. 271, Div. of Timber Mgt. Res., U.S.D.A., Washington, D.C. 762 pp.
HOLLAND, W. J. 1903. The moth book, Doubleday, Page & Co., New York. 479 pp.
VILLIARD, P. 1969. Moths and how to rear them. Funk & Wagnalls, New York.

242 pp.

Journal of the Lepidopterists’ Society
33(3), 1979, 166

DOUBLY OVERWINTERING CITHERONIA REGALIS FABRICIUS
(LEPIDOPTERA: SATURNIIDAE)

In 1975 I reared 94 larvae of Citheronia regalis Fabricius which overwintered suc-
cessfully. Among these were 50 males and 44 females. In 1976, many eclosed, but 18
males and 5 females remained dormant. This represents 36% of the males and 10% of
the females, or 24.5% of the total sample.

Later, in 1977 I secured two matings between doubly overwintered moths. From ova
of those females, I reared 41 larvae, the pupae of which were still viable in the spring
of 1978. I could now observe whether or not the tendency of these pupae to remain in
diapause over their first summer had been reinforced by the selective matings of their
doubly-overwintered parents.

The pupae consisted of 20 males and 21 females. Five of these died before emer-
gence. Among these, two males and one female developed but failed to eclose, while
one male and one female showed no signs of metamorphosis. The five dead pupae
were excluded from the data.

Of the remaining 17 males, 15 emerged in 1978, with only two (11.8%) remaining in
diapause. Of the 19 females, 16 emerged, and only three (15.8%) remained in diapause.
Thus of the total population 13.9% remained in diapause.

These observations do not support the hypothesis that double overwintering is under
simple genetic control. I could not discern any other determining conditions for the
phenomenon, though double overwintering must obviously have survival value for the
species by carrying it through occasional catastrophic years.

C. BROOKE WorTH, Eldora, Cape May Co., R. D. Delmont, New Jersey 08314.

Journal of the Lepidopterists’ Society
33(3), 1979, 167-169

A DOCUMENTATION OF BIENNIALISM IN
BOLORIA POLARIS (NYMPHALIDAE)!

JOHN H. MASTERS
25711 North Vista Fairways Drive, Valencia, California 91355

ABSTRACT. Capture/noncapture records of Boloria polaris at Churchill, Manitoba
over a 46-year timespan provide documentation that the species (at least at Churchill)
is biennial, where adults fly only in odd-numbered years.

Biennialism in insects is that situation where an insect’s life-cycle
takes two years to complete so that imagos are produced only after
two years of pre-imaginal development. It may be accompanied by
biennial flights when, in a given locality, adults fly only in alternate
years (periodical flight); or it may be accompanied by annual flights.
Unless biennial flights are involved, biennialism is extremely difficult
to perceive in nature without carefully working out the insect’s life
history. Annual flights may occur when the species is only partially
biennial or when two allochronic populations are involved. Docu-
mented cases of regular biennialism are very rare in Lepidoptera and
heretofore have been confined to two species of moth (Lasiocampa
quercus callunae in Europe and Coloradia pandora lindseyi Barnes
and Benjamin in North America) and a few species of Satyridae (e.g.,
Oeneis macounii Edwards, Oeneis nevadensis Felder and Felder,
Oeneis jutta Hubner, Erebia claudina Borkhausen, and Erebia ligea
Linnaeus). For a review of periodical flight behavior in other insects,
see Bulmer (1977).

In working on a study of butterflies occurring at Churchill, Mani-
toba (Masters, 1971), it became evident to me that Boloria polaris
stellata (Masters) is biennial at Churchill and since that time, I have
accumulated enough data to document this fact rather well. This is
the first documentation of periodical biennialism in Nymphalid but-
terflies. It has been suggested, and recent collecting data tends to
support the contention, that several other species of Boloria (e.g., B.
distincta Gibson and B. chariclea Schneider) and the arctic popula-
tions of one species of Hesperiidae (Hesperia manitoba borealis
Lindsey) will also turn out to be biennial.

Churchill, Manitoba, since the opening of the Hudson Bay Railroad
in 1930, has been a classic arctic collecting locality. Because Churchill
has been collected repeatedly for a span of 46 years, it is possible to
document the capture and noncapture data for Boloria polaris from

1 Originally submitted and accepted for publication in 1971; publication delayed due to manuscript loss in the
mail. Revised to incorporate new data.

168 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

TABLE 1. Capture/noncapture records of Boloria polaris at Churchill, Manitoba.

Expeditions recording Expeditions failing to
Boloria polaris record Boloria polaris

1932 (A. V. Harper)

1936 (H. E. McClure)

1937 (G. S. Brooks)

1939 (G. S. Brooks)

1939 (B. Wilk) 1940 (G. S. Brooks)

1941 (G. S. Brooks) 1942 (G. S. Brooks)

1943 (G. S. Brooks) 1944 (G. S. Brooks)
1946 (G. S. Brooks)

1947 (T. H. Freeman)

1951 (A. B. Klots & R. D. Bird) 1952 (A. B. Klots & R. D. Bird)

1961 (F. H. & P. W. Chermock)

1963 (F. H. & P. W. Chermock)

1967 (J. A. Ebner)! 1968 (C. S. Quelch)

1969 (A. E. Brower) 1969 (J. H. Masters)?
1970 (J. H. Masters)
1970 (C. McCullough)?

1971 (C. D. Ferris)
1974 (D. Oosting & D. Parshall)
1976 (D. Oosting & D. Parshall)
1978 (J. Troubridge)

1 Specimens presumably not taken by Ebner personally.
* Expeditions in late July and August, too late in the year to expect to find Boloria polaris.

27 collecting expeditions. These data are tabulated in Table 1. Infor-
mation was gleaned from 14 expeditions to Churchill in odd-num-
bered years; 13 of which recorded B. polaris (only my very short trip
in late July 1969 did not). On the other hand, I have documented 13
expeditions to Churchill in even-numbered years, none of which re-
corded B. polaris, even though 12 of them were at the proper time of
year. I have also scanned both private and institutional collections for
specimens of B. polaris that might have been collected at Churchill
in even-numbered years, and have failed to find a single specimen.
The evidence is thus conclusive, albeit circumstantial, that Boloria
polaris has an odd-year only biennial flight at Churchill.

Additional records of Boloria polaris accumulated from other lo-
calities suggest that it is probably biennial everywhere, but may be
subject to random geographic alternation of odd-year and even-year
broods. These data are summarized in Table 2. Unfortunately, no
other region in the arctic has been collected as extensively as Chur-
chill for butterflies, over so long a period, and a comparable docu-
mentation of the biennial flight of B. polaris at any additional locality
cannot be made. The collecting reports of the Alaska Lepidoptera

VOLUME 33, NUMBER 3 169

TABLE 2. Circumpolar capture records of Boloria polaris, noting year of capture.
Records, other than Churchill, based on specimens in the American Museum of Natural
History, New York and Camegie Museum, Pittsburgh.

Locality Odd-year captures Even-year captures
Norway: Maalselvin 1923, 1935, 1937,
1939,1971
Finland 1934, 1936, 1946
Alaska:
Eagle Summit & Mt. 1931, 1933, 1955, 1961,
McKinley 1969, 1971
Nome 1968
Yukon: Haines Junction 1966, 1968
British Columbia: Atlin
& Summit Lake 1930, 1966
Mackenzie: Mackenzie
Delta & Coppermine 1942, 1966
Keewatin: Baker Lake
& Eskimo Point 1952, 1956, 1966, 1968
Manitoba: Churchill 1933, 1937, 1939, 1941,
1943, 1947, 1951, 1961,
1963, 1967, 1969, 1971
Greenland 1957! 1922, 1926, 1932, 1958
Baffin Island 1925, 1934 1896, 1954

1 One specimen 22 July 1957 by A. T. Washburn.

Survey, over a ten year period (1967-1977) establish reasonably well,
however, the biennial flight of the species on Ester and Murphy
Domes near Fairbanks, Alaska.

LITERATURE CITED

BULMER, M. G. 1977. Periodical insects. Am. Nat. 111: 1099-1117.
MASTERS, J. H. 1971. The butterflies of Churchill, Manitoba. Mid-Continent Lepid.
Ser. 25: 1-13.

Journal of the Lepidopterists’ Society
33(3), 1979, 170-173

FOODPLANT OF ALPINE EUPHYDRYAS ANICIA
(NYMPHALIDAE)

RAYMOND R. WHITE
Dept. of Biological Sciences, Old Dominion University, Norfolk, Virginia 23508

ABSTRACT. Alpine Euphydryas anicia use Besseya alpina as their foodplant
during pre- and post-diapause development.

Until 1974 the foodplant of alpine Euphydryas anicia Doubleday
& Hewitson was unknown. Discovery of the use of Besseya alpina
Rydberg (Scrophulariaceae) by a population at Cumberland Pass
(Gunnison Co., Colorado) was mentioned by Ehrlich et al. (1975). My
purpose here is to document the more widespread use of this plant
by alpine E. anicia and to elicit observations by other workers in the
field.

Figure 1 shows the distribution of alpine populations investigated
to date. Table 1 shows evidence for the use of B. alpina by these
populations. At the present time two things are known about alpine
E. anicia populations: first, they occur on almost every peak above
3,660 m elevation above sea level in Colorado; second, the oviposition
plant and post-diapause larval foodplant is B. alpina for all popula-
tions studied.

Unfortunately, we know nothing of the factors that limit the sizes
of populations of alpine E. anicia, though we have learned something
of their movement patterns (Cullenward et al., 1979). Since slow-melt-
ing snowbanks make early access to the alpine very difficult and since
larvae are difficult to find, few post-diapause larvae have been col-
lected. A few parasitoids have been reared from the 70 larvae col-
lected to date. These are currently being identified. Few observations
have been made on these larval populations in August and none in
September, when predation and starvation of pre-diapause larvae
might be seen. One can, however, infer something about the plant-
herbivore relationship by observing the distribution and habit of plant
growth and the magnitude of post-diapause feeding damage (Table 1).

Bessya alpina is perennial and strictly alpine, known from 3,500 to
4,350 m in Wyoming, Utah, Colorado and New Mexico. The distri-
bution often includes points of greatest local elevation (sites 1, 3, 11,
and 12), but often does not, apparently due to lack of soil just beneath
the scree at some of these points. Often the plants grow right up to
edge of exposed ridges and sometimes on the ridge tops themselves.

In general, the sparse populations of Besseya that are often pros-
trate in form and hidden among rocks suffer less feeding damage than

VOLUME 33, NUMBER 3 ol

ee

Oger 7

12 6 5
GUNNISON

Fic. 1. Map of Colorado showing peaks investigated. Names on map refer to coun-
ties. 1, Mt. Evans; 2, Loveland Pass; 3, Cottonwood Pass; 4, Cumberland Pass; 5,
Double Top; 6, Crested Butte; 7, Gothic Mtn.; 8, Avery Peak; 9, Mt. Bellview; 10,
Mt. Baldy; 11, Cinnamon Mtn.; 12, Ruby Peak; 13, Rock Ridge.

172 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

TABLE 1. Distribution and habit of Besseya growth, with post-diapause feeding
damage. Locality numbers refer to Fig. 1 map.

Proportion

of crop Number Number
consumed by of egg _ of post— Number
post-diapause masses diapause of plants Plant Food
Locality larvae found larvae sampled Plant density habit plant?
1 0.002 0) 0) 125 very sparse Ih No
D2 0.007 0) ®) 299 sparse 1 No
3 — z 0 50 sparse 2 Yes
4 0.09 23 15 1,000 dense in 3 Yes
places
o 0.01 0 0 130 very sparse I No
6 0.15 0 0 6 very sparse i No
7 0.04 3 IL 130 sparse 2 Yes
8 0.02 ®) ®) 172 common in 3 No
places
9 0.08 2 1 102 common 3 Yes
10 0.06 1 2D, 114 common in 3 Yes
places
ll 0.09 5 3 100 common over a 3 Yes
small area
12 0.09 3 1 ae, common 3 Yes
13 0.06 4 2 132 common Dip) Yes

Plant habits: 1—among rocks, small and prostrate; 2—in scree, but not inconspicuous; 3—in the open, erect.

do the denser, more conspicuous populations. Three alternative hy-
potheses may explain this phenomenon: first, some environmental
variable may curtail both plant and butterfly growth; second, the but-
terflies may not be able to generate large populations where the plants
are scarce due to some factor other than the butterfly itself; third, the
butterflies may cause the observed pattern by over-grazing conspic-
uous populations of Besseya. Further work is planned to learn which
of these hypotheses is correct.

The magnitude of pre-diapause larval feeding damage and therefore
its importance to the Besseya populations remains unknown. The
magnitude, however, is likely to exceed substantially that of post-
diapause feeding damage (Table 1). The eggs are laid in clusters of
90-100, so a large number of larvae feed on a single plant. The larvae
must reach a diapause weight of 5-10 mg each. Since the conversion
rate of Besseya biomass to larval biomass is probably about ten to one
(White, 1973), each brood may consume 2.5-10 g [(50-100) x (5-10) x
(10)| of Besseya. This amount, according to a small sample of Besseya
leaves, is comparable to the size of the average Besseya plant (2.7-
9.5 g, varying by year and by population). Pre-diapause larval growth
appears to be so slow at these elevations that significant plant growth

VOLUME 33, NUMBER 3 173

might occur during the feeding period. Thus, pre-diapause feeding
might usually be completed on a single plant without larval migration
or its consequent starvation. Since it appears that Besseya plants
either bloom and set seed early in a growth season or not at all that
season, larval feeding during the growing season does not affect seed
set in that year. Sufficient defoliation in one year may suppress seed
set in the future or may cause death of the plant. About 20% of the
plants set seed in a given year. Presumably this proportion varies with
a number of factors, including the extent of defoliation.

One frequently finds significant numbers of both sexes of E. anicia
flying 100-300 m below the lowest part of Besseya distribution (sites
4,5, 7, 8, and 12) and sometimes some distance away. Mark-release-
recapture work at Cumberland Pass (site 4) showed that at least sev-
eral hundred individuals fly in, and tend to stay in, an area several
hundred meters from the nearest Besseya. At Double Top (site 5)
numerous individuals occupy large areas apparently devoid of Bes-
seya. Also, at some alpine sites (1, 2, 5 and 8 of Fig. 1) where E. anicia
is known to maintain populations, no egg masses were found on Bes-
seya and feeding damage was light. Why these things should be so if
these populations are utilizing Besseya and represent the same eco-
type as do the others remains mysterious. One can only suspect that
superficially similar local populations are in fact ecologically rather
heterogeneous.

ACKNOWLEDGMENTS

I thank Dr. Paul R. Ehrlich for his encouragement and Dianne
Stewart for technical assistance. This work was partially supported by
a grant from the Old Dominion University Research Foundation and
was carried out from a base at the Rocky Mountain Biological Labo-
ratory in Gothic, Colorado.

LITERATURE CITED

CULLENWARD, M. J., P. R. EHRLICH, R. R. WHITE & C. E. HOLDREN. 1979. The
ecology and population genetics of an alpine checkerspot butterfly, Euphydryas
anicia. Oecologia 38: 1-12.

EHRLICH, P. R., R. R. WHITE, M. C. SINGER, S. W. MCKECHNIE & L. E. GILBERT.
1975. Checkerspot butterflies: An historical perspective. Science 188: 221-228.

WHITE, R. R. 1973. Community Relationships of the Butterfly, Euphydryas editha.
Ph.D. dissertation, Stanford University.

Journal of the Lepidopterists’ Society
33(3), 1979, 174-178

FOURTH ADDITION TO THE SUPPLEMENTAL LIST OF
MACROLEPIDOPTERA OF NEW JERSEY

JOSEPH MULLER
R. D. 1, Lebanon, New Jersey 08833

ABSTRACT. Additional species, subspecies and named aberrations (with larval
food plant, when known) are added to the list of Macrolepidoptera of New Jersey not
recorded in John B. Smith’s Report of the New Jersey State Museum (1909) or in my
later reports to date (1965, 1968, 1973, 1976).

The following new additions were collected by the author, Dr. C.
Brooke Worth, Dr. Dale Schweitzer, and the Department of Agricul-
ture at Rutgers University. References are cited for those species al-
ready reported elsewhere in the literature.

In this area, light pollution appears to be the main reason for the
decrease in moths. The only collecting locality known to me where
lights and bait still work is Dr. Worth’s farm at Eldora, (Cape May
Co.). This area is relatively isolated, since the land mass is on a pen-
insula. Most sphingids, saturniids and other conspicuous Lepidoptera
formerly common throughout the state are still found here in large
numbers.

Checklist numbers and classification for moths are taken from
McDunnough (1938), updated to current nomenclature. Localities,
dates of collection, collector (and when known, foodplant) are shown.
Specimens not followed by the name of the collector were taken by
the author. Specimens collected at Eldora, Cape May Co., were col-
lected by the author in collaboration with Dr. Worth.

ACKNOWLEDGMENTS

I wish to thank Dr. Frederick Rindge and Eric Quinter, of the
American Museum of Natural History, for determining some of the
following specimens and for reading the manuscript, and Dr. Rindge
for providing references to published records.

MOTHS
Arctiidae

Holomelina Herrich-Schaeffer
1022 opella form ““‘belmaria’” Ehrman
Eldora, 17 June; 17 Sept.
Dandelion, Taraxacum officinale; plantain, Plantago major

VOLUME 33, NUMBER 3 IGS

Noctuidae
Noctuinae
Graphiphora Ochsenheimer
1525 tenuicula Morrison
Drakestown, 17 Sept., Rutgers
Hadeninae
Leucania Ochsenheimer
1978-1 linda Franclemont
Long Valley, 21 July, Rutgers
Cuculliinae

Metaxaglaea Franclemont
2221-2 ni: sp.
Eldora, 16 Oct., 25 Nov., 1, 11 Jan.
American holly, Ilex opaca

Amphipyrinae
Apamea Ochsenheimer
2328-1 inebriata Ferguson
Lakehurst, 8 April, 29 June, 25 July, Lemmer; Jerseyville, 9 July, Shulgin; Free-
hold, 17 July, Franclemont
Agroperina Hampson
2373 lutosa Andrews
Woodglen, 25 April, 2 July, Rutgers
Luperina Boisduval
2393 passer Guenée form “conspicua’” Morrison
Drakestown, 21 Sept., Rutgers
Dock, Rumex occidentalis
Oligia Hubner
2421 minuscula Morrison
Lakehurst, 6 Sept., 15 Oct., C. Rummel
Xylomoia Staudinger
2434 chagnoni Barnes & McDunnough
Seabrook, 12 June, Rutgers
Borer in Phalaris arundinacea
Callopistria Hubner
2538 floridensis Guenée
East Millstone, 27 Sept., Rutgers
Ferns
Platyperigea Smith
2655 multifera Walker
Freehold, 25 June, Rutgers
2656 extima Walker
Freehold, 25 June, Rutgers
Bellura Walker
2709 melanopyga Grote
Camden, 25 May; Delanco, 26 May, Rutgers
Borer in yellow water lily

176 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

Acontiinae

Cryphia Hubner
3100 villificans Barnes & McDunnough
Oliver, 12 June, Rutgers
Tarachidia Hampson
3185 semiflava Guenée
Eldora, 20 May

Catocalinae

Catocala Schrank
3336 paleogama Guenée, form “denussa” Ehrman
Hunterdon Co., 8 Sept., 9 Oct.
Hickory, Carya; walnut, Juglans
3386 gracilis Edwards, form “lemmeri’” Mayfield, 1923
Lakehurst, 1 Sept., Lemmer
Vaccinium
3388 herodias Barnes & Benjamin, Gerhardi Benjamin & Barnes
Lakehurst, 15 July, Rutgers
3410 micronympha Guenée, form “lolita” Sargent
Lebanon, ex ovo 3, 8, 18, 20, 22 June, 3 Sept.
Red oak, Quercus rubra
Zale Hubner
3493 metata Smith
Eldora, 20 May
Pinus virginiana

Ophiderinae
Metalectra Hubner
3658-1 richardsi Brower
Eldora, 23 July
Hypeninae

Hypenodes Doubleday
3728-1 fractilinea Smith
Martinsville, 10 June, Rutgers
Parahypenodes Barnes & McDunnough
3729 quadralis Bames & McDunnough
Hughesville, 25 July, Rutgers

Herminiinae
Epizeuxis Hubner
3735 concisa Walker
Woodglen, 25 July; Evesboro, 9 July, Rutgers

Notodontidae

Heterocampa Doubleday
3890 varia Walker
Chatsworth, 12 July, J. Madenjian
Oak, Quercus sp.

VOLUME 33, NUMBER 3 177

Geometridae
Larentiinae

Coryphista Hulst
4248 meadii atlantica Munroe, 1954
Irvington, May—Sept., E. E. Witte
Barberry
Eupithecia Curtis
4266-1 slossonata McDunnough, 1949
Eldora, 17 May
Hydriomena Hubner
4477 transfigurata Swett. McDunnough, 1954
Lebanon State Forest, 13 June, Rutgers; Lebanon, 7, 13, 25 May

Ennominae

Lomographa Hubner
4608 glomeraria Grote
Lebanon, 16 May
Hypomecis Hubner
4738 gnopharia (Guenée)—formerly in Pseudoboarmia. Rindge, 1973
Eldora, 25 May, 18 June
Anacamptodes McDunnough
4917-1 humaria humaria (Guenée). Rindge, 1966
Eldora, 20 May
General feeder on deciduous plants
Glenoides McDunnough
4948 texanaria Hulst
Eldora, 24 Oct.
Epimecis Hubner
4951 virginaria Cramer, form “dendaria’ Guenée
Eldora, 20 May
Tulip tree, sassafras
Philgalia Duponchel
4956 strigataria (Minot). “Melanic male.” Rindge, 1975
Lebanon, 5 April
4957 denticulata Hulst. Rindge, 1975
Orange Mts., Essex Co., 3 March, Lemmer. In Cambridge collection.
Tacparia Walker
0024 atropunctata (Packard). Ferguson, “1973” (1974)
Near Lisbon, 12 June, J. Madenjian
Birch, Betula sp.
Sicya Guenée
5161 macularia Harrison
Lebanon, 17 July
Spirea, Ceanothus

LITERATURE CITED

FERGUSON. “1973” (1974) Proc. Entomol. Soc. Wash. 75: 476. (Lists two N.J. locali-
ties.)

FERNALD, M. L. 1950. Gray’s New Manual of Botany, Eighth Ed.

FORBES, W. T. M. 1923-1960. Lepidoptera of New York and Neighboring States, Parts
II, II] and IV. New York Agr. Exp. Sta. at Cornell, Ithaca, New York.

MAYFIELD, T. D. 1923. Bull. Brooklyn Entomol. Soc. 18: 33.

178 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

McCDUNNOUGH. 1949. Bull. Amer. Mus. Nat. Hist. 93: 548. (Cites Lakehurst, N.J.)

1954. Bull. Amer. Mus. Nat. Hist., 104: 279. (Cites Lakehurst, N.J.)

MULLER, J. 1965. Supplemental List of Macrolepidoptera of New Jersey. J. New York
Entomol. Soc. 73: 63-77.

1968. Additions to the Supplemental List of New Jersey Macrolepidoptera. J.

New York Entomol. Soc. 76: 303-306.

1973. Second Addition to the Supplemental List of Macrolepidoptera of New

Jersey. J. New York Entomol. Soc. 81: 66-71.

1976. Third Addition to the Supplemental List of Macrolepidoptera of New
Jersey. J. New York Entomol. Soc. 84: 197-200.

MUNROE. 1954. Canadian Entomol. 86: 282. (Type locality: Irvington, New Jersey).

RINDGE, F. 1966. Bull. Amer. Mus. Nat. Hist. 132: 216. (Fig. 7 [distribution map]; six
localities from New Jersey).

1973. Amer. Mus. Novitates No. 2514: 23. (Fig. 24 [distribution map]; five

localities for New Jersey).

1975. Bull. Amer. Mus. Nat. Hist. 156: 124. (Fig. 76 [distribution map]; many

New Jersey localities).

1975. idem. p. 128. (Fig. 76 [distribution map]; many New Jersey localities.)

SMITH, J. B. 1910. Report of New Jersey State Museum for 1909. Trenton.

Errata

Corrections in Third Addition to the Supplemental List of Macrolepidoptera of New
Jersey (1976):

Page 197. No. 248. polixenes should read polyxenes.
Page 197. Nyphalidae should read Nymphalidae.

Page 198. Meliana Curtis = Leucania Ochsenheimer.
Page 199. Stabidium Grote = Plagiomimicus Grote.
Page 199. Camptylochia Stephens = Epizeuxis Hubner.
Page 199. 3777 should read 3735.

Page 199. Epicnaptera Rambur = Phyllodesma Hubner.
Page 200. Zanolidae = Apatelodidae.

Page 200. No. 4932. Mistaken identity.

Journal of the Lepidopterists’ Society
33(3), 1979, 179-188

NEW STATUS FOR EPIBLEMA MINUTANA (KEARFOTT) AND
NEW SPECIES OF EPIBLEMA HUBNER AND
SONIA HEINRICH (TORTRICIDAE)

A. BLANCHARD
P. O. Box 20304, Houston, Texas 77025

ABSTRACT. Epiblema minutana is a distinct species, differing from E. strenuana
(of which Heinrich considéred it a synonym) in size, wing shape, male and female
genitalia. Epiblema luctuosana is described from southern and eastern Texas. Sonia
paraplesiana n. sp., widely distributed in the eastern United States, has long been

confused with Zellers Sonia constrictana known only from south Texas and south
Florida.

Epiblema minutana (Kearfott) revised status

Eucosma minutana Kearfott 1905, Proc. U.S.N.M. 28: 356.
Eucosma antaxia Meyrick 1920, Exot. Microlepid. 2(2): 344.
Epiblema strenuana (Walker 1863) Heinrich 1923, U.S.N.M. Bull 123: 140.

Remarks. My interest in this problem was arouséd when my wife and I took on
North Padre Island, Texas, in March and again in June 1978, a series of specimens of
an Epiblema which in Heinrich’s key (1923, p. 137) keyed out at strenuana, but which,
after examination of the female genitalia proved to be definitely different.

Kearfott’s description of E. minutana states that minutana is of very much smaller
size than strenuana and that the forewings are much narrower than strenuana—more
than three times as long as wide. As our Padre Island specimens presented these two
characters, I requested help of the National Museum of Natural History and the Amer-
ican Museum of Natural History which have most of Kearfott’s types of E. minutana.

Kearfott described minutana from a series of about 40 specimens, 15 of which are
before me: 9 from NMNH and 6 from AMNH. I eliminate from consideration one of
the specimens borrowed from NMNH because, although labelled type by Kearfott, it
does not appear to be conspecific with the other fourteen.

These fourteen types made a remarkably homogeneous group and my small, narrow-
winged specimens from Padre Island, as well as some of my smallest “strenuana” go
very well with them. Figs. 2, 4, 6, 9, and 10 permit easy comparison of Kearfott’s types
with some recently collected specimens. I do not include here a list of the Texas
material examined in this study.

Description of female. Table 1 summarizes data for 22 dissected females. Among
the females two characters are practically diagnostic: (i) the diameter at base of the
signa, and (ii) the shape of the lamella postvaginalis. The signa are two hollow, curved,
thormlike, sclerotized processes flattened at their distal end, round at their base. It is
almost always easy to accurately measure their base diameter. Signa of both groups of
specimens are about the same size, but they average larger (with no overlap in size)
in strenuana than in minutiana. The lamella postvaginalis is also consistently different.
In strenuana it is longer in proportion to width and its sides are straight and parallel;
in minutana the sides diverge caudad. Table 1 shows that there is overlap in fore-
wing length as well as in forewing length-to-width ratio. The corpus bursae size and
shape are not very good characters either, although the minutana corpus bursae may be
more pear-shaped, less bulbous than that of strenuana.

Description of male. There appears to be little difference between males and
females regarding wing size and forewing length-to-width ratio (Table 2). Some males
will prove difficult to identify: size and length-to-width ratio are good characters if they
do not fall in the range of overlap. When both these characters fail, one may rely on
the size of genitalia or the shape of the valves: the neck incurvation of the valves of
minutana is usually shallower than that of strenuana.

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

180

Fics. 1-6: Epiblema minutana. 1, 3 lectotype; 2, 6 specimen from No. Padre
Island, Nueces Co., 27 July 1978; 3, genitalia of lectotype, slide U.S.N.M. 24505; 4,
genitalia of specimen of Fig. 2, slide A.B. 4473; 5, 2 genitalia of lectoparatype, Mont-
clair, New Jersey, July 8, slide U.S.N.M. 24501; 6, genitalia of a 2 specimen from No.
Padre Island, 8 July 1978, slide A.B. 4507.

VOLUME 33, NUMBER 3 181

TABLE 1. Comparison of measurements on dissected female specimens of Epiblema
minutana and E. strenuana. Measurements are given as the median (and range).

Forewin
Forewing feeicehvaiach Corpus burae Sigma base
Species length! ratio length: width! diameter!
Epiblema minutana, 5.6 Sell 1.15; 0.85 0.08
11 specimens (4.7-6.3) (2.8-3.35) (0:95; O.5=1745: 1.0) ~~ (0:07—0.11)
Epiblema strenuana, 7.8 2203 AS le 0.14
11 specimens (9:6=820) . (2-5-2959) GEO Se Sle) (ORE2—0. 18)

1 Measurements given in millimeters.

Foodplant. The foodplant of E. minutana, according to Forbes (1923: 413) is Am-
brosia artesmisiifolia and I am convinced that, when better known, E. minutana will
prove to be as widely distributed as its foodplant.

Type data. Lectotype, hereby designated, 6, Montclair, New Jersey, 8 July. Geni-
talia on slide USNM 24505 (Figs. 1, 3 & 7). Lectoparatypes: 2, Montclair, New Jersey,
8 July; left pair of wings and genitalia on slide USNM 24501 (Figs. 5, 8). 6, Tryon,
North Carolina, 23 May 1904, Fiske collector, right pair of wings and genitalia on slide
USNM 24506. ¢, Tryon, North Carolina, 24 May 1904, Fiske collector, the metathorax
and hind wings have, somewhat improperly, been glued. ¢, Cincinnati, Ohio, 10 July
1904, Annette Braun, abdomen missing. 2, Plummer’s I., Maryland, July, Aug. Busck,
abdomen and antennae missing. 6, New Brighton, Pennsylvania, 20 May 1903, H. D.
Merrick, abdomen and some of both antennae missing. 2, 440 Liss. 11 Jan. 1901, LHW
missing. All of these are in NMNH. 6, Essex County Park, New Jersey, 13 Aug.,

TABLE 2. Forewing measurements on fourteen of Kearfott’s types of Epiblema mi-
nutana.

Type specimen Forewing
Length! Length-to-width
Museum Sex & number (mm) ratio?
NMNH 6 1(USNM 24505) 4.9 oe
6 3(USNM 24506) 5.6 SID)
3 4 6.1 SO)
6 9 DD Oro)
Gil 6.1 SD)
AMNH ul oh Or
3 2 DO SY
6 e) 4.9 2.95
6 9 5.4 2.8
NMNH Q 2 5.4 3.45
OIG 0.6 oD
28 6.1 one
AMNH 24 5D Si5D)
26 5.4 SD

1 Mean for males—5.55 m; for females—5.6 mm
2 Mean for males—3.25; for females—3.30

182 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

Fics. 7-10, 16,17: Epiblema venation. Figs. 7-10: E. minutana, 7, 3 lectoparatype,
slide U.S.N.M. 24506; 8, 2 lectoparatype, slide U.S.N.M. 24501; 9, 3 from No. Padre
Island, 6 April 1978, slide A.B. 4472; 10, 2 from same date and location, slide A.B.
4465. Figs. 16, 17: E. luctuosana. 16, 3 paratype from Padre Island National Seashore,
24 June 1976, slide A.B. 4269; 17, 2 paratype of Fig. 15, slide A.B. 4521.

VOLUME 33, NUMBER 3 183

Fics. 11-15: Epiblema luctuosana. 11, 6 holotype; 12, 2 paratype, No. Padre Is-
land, 9 June 1978; 13, 3 paratype, No. Padre Island, 5 June 1978; 14, genitalia of
holotype, slide A.B. 4468; 15, genitalia of a 2 paratype, No. Padre Island, 5 June 1978,
slide A.B. 4521.

trap, W. D. Kearfott, in good condition, No. 34716. 3, Essex County Park, New
Jersey, 20 Aug., trap, W. D. Kearfott, head missing, otherwise in good condition,
No. 34716. Essex County Park, New Jersey, 20 July, trap, W. D. Kearfott, abdomen
missing, No. 34716, Female, Wilkesbarre, Pennsylvania, 6 June, W. D. Kearfott, geni-
talia on slide A.B. 4598. 3, Montclair, New Jersey, u. m. s. 10 May, W. D. Kearfott,
mildewed, Ac. 4667. ?, Cincinnati, O., 18 June 1904, Annette Braun, traces of mildew,
Ac. 4667. All six lectoparatypes are in AMNH.

Some material collected by A. & M. E. Blanchard, discussed above will be deposited
in NMNH and AMNH. All the females collected by A. & M. E. Blanchard, described
in Table 1 will (together with their genitalia slides) be deposited in NMNH.

I have been advised by a reviewer of this article that Richard L. Brown (1973:58) in
his unpublished Masters thesis had already listed E. minutana as a species. It is a
pleasure to give him credit here.

184 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

Epiblema luctuosana A. Blanchard, n. sp.

Description. Palpi exceeding front by a little less than an eye diameter, third seg-
ment smoothly scaled, almost completely hidden in the loose, grayish scaling of the
much longer second segment. Antennae simple, ciliate beneath, the sensory hairs hard-
ly exceeding the scales. Male and female venation is shown in Figs. 16 and 17; mac-
ulation shown in Figs. 11, 12 and 13. Nearly all the scales, except the whitish ones of
the ocelloid patch and those of the whitish patch at the middle of the dorsum are black
or blackish basally, white or whitish distally, without appreciable transition: this is
true, not only of the wing and body scales, but even of those which clothe the palpi,
the antennae and the legs. It gives this insect a pepper-and-salt appearance which is
most noticeable under the microscope at low magnification. The size and appearance
of the ocelloid patch varies little but the whitish patch near the middle of the dorsum
varies considerably, from very large (Fig. 13) to almost obsolete (Fig. 11), to completely
obsolete. Hind wing dark fuscous.

Wing expanse: males 9.3-13.5 , mean = 12 mm; females 11.8-13.6 mm, mean =
12.7 mm.

Male genitalia (Fig. 14). From the holotype, slide A.B. 4468.

Female genitalia (Fig. 15). Slide A. B. 4521, paratype from North Padre Island,
Nueces Co., Texas, 5 June 1978. Ductus seminalis from ductus bursae somewhat closer
to genital opening than to corpus bursae. Signa very small, conical.

Type data. Holotype (Fig. 11): 6, North Padre Island, Nueces Co., Texas, 6 April
1978, genitalia on slide A.B. 4468 (Fig. 14), deposited in NMNH (type No. 75822).
Paratypes: (All localities in Texas) 3 2 2, Houston, 5-6 June 1968; 2, Deutschburg,
Jackson Co., 2 Aug. 1972; Santa Ana Refuge, 27 Nov. 1973, 6; 4 Nov. 1974, 6; Sinton,
Welder Refuge, San Patricio Co.; 30 June 1975, 2; North Padre Island, either Nueces
Co., or National Seashore, Kleberg Co., 22 April 1976, 6; 19 May 1976, 56, 4 29; 24
June 1976, 2 66; 19 July 1976, 5; 20 May 1977, 6; 18-21 June 1977, 3 66,3 22;6
April 1978, 2 646; 5-10 June 1978, 16 63,9 22; 6, Conroe, Camp Strake, Montgomery
Co., 14 Sept. 1977; 2, Eagle Lake, Attwater Prairie Chicken Refuge, Colorado Co. The
paratypes are in the author's collection.

I am indebted to Richard L. Brown for examining some of my specimens and slides
and I quote him (in litt.) regarding the relationships of luctuosana to other Epiblema
species: “E. luctuosana appears to be related with the species numerosana (Zeller),
praesumptiosa Heinrich, grossbecki Heinrich, abruptana (Walsingham) and deflexana
Heinrich. This group of species lacks sclerotization of the female ductus bursae (female
of deflexana not examined) and has longer coronal setae on the cucullus relative to
other Epiblema species. This group is also characterized by the vinculum depth ex-
ceeding the tegumen length and the presence of black scales on the labial palpi. E.
luctuosana shares the above characters with this species group, but is distinguished
by the absence of black scales on the labial palpi.”

Sonia paraplesiana A. Blanchard, n. sp.

Head. Front clay yellow; vertex pale brown; first and second segments of palpi shaggily
squamous underneath, with a laterally compressed brush of long, clay yellow, brown
dotted scales; third segment smoothly scaled, more than half hidden in scaling of
second segment, brown with clay yellow tip. Antennae simple, shortly pubescent,
brownish.

Maculation. As in Fig. 18, the color from dark fuscous brown to similarly tinted
whitish. Wing venation: Figs. 24 and 25; the length of the stalk of veins R, and M, of
the hind wing varies from short to about half as long as their free parts, but these two
veins appear to be consistently stalked.

Wing expanse. 12.5-17 mm.

Male genitalia. As in Fig. 20.

Female genitalia. As in Figs. 22 and 26.

Type data. Holotype: 3, Houston, Texas, 5 June 1968, genitalia on slide A.B. 4365,

VOLUME 33, NUMBER 3 185

Fics. 18-23: Sonia species. Figs. 18, 20, 22: S. paraplesiana. 18, holotype; 20, ¢
genitalia of a paratype, Houston, Texas, 8 June 1968, slide A.B. 4524; 22, 2 genitalia
of a paratype, Sinton, Texas, Welder Wildlife Refuge, 2 June 1978, slide A.B. 4512.
Figs. 19, 21, 23: S. constrictana. 19, 6 from Welder Refuge, 30 June 1975; 21, 6
genitalia of a specimen from Santa Ana National Refuge, Texas, 15 September 1974,

slide A.B. 4523; 23, 2 genitalia of a specimen from Welder Refuge 30 June 1975, slide
A.B. 4520.

186 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

Fics. 24-29: Sonia species. Figs. 24, 25: S. paraplesiana venation. 24, d paratype,
slide A.B. 0892; 25, 2 paratype, slide A.B. 4512. Figs. 27, 28: S. constrictana vena-
tion. 27, 6, slide A.B. 4283; 28, 2, slide A.B. 4358. Figs. 26, 29: lamella
postvaginalis: 26, S. paraplesiana, 2 paratype, slide A.B. 4512; 29, S. constrictana,
2, slide A.B. 4360.

deposited in NMNH, No. 76098. Paratypes: Houston, Texas, 21 June 1966, 6; 26
August 1966, 6; 7 November 1966, 6; 4, 5 June 1967, 2 36; 5-8 June 1968, 7 36d,
?; %, Sinton, Welder Wildlife Refuge, San Patricio Co., Texas, 30 June 1975; 2, Town
Bluff, Tyler Co., Texas, 15 Sept. 1975; 2 66, Tennessee Colony, Anderson Co., Texas

ee 1978; A. & M. E. Blanchard collectors. Specimens presently are in the author's
collection.

VOLUME 33, NUMBER 3 187

32

Fics. 30-32: Sonia constrictana. 30, 3 holotype; 31, genitalia, slide V. Adams
XI1.4.69; 32, venation of right pair of wings, same slide.

The following paratypes are in the National Museum: Putnam Co., Illinois, 26 June
1964, 3; 2 July 1964, ¢; 1 July 1965, 6; 25 June 1966, 6; 7 July 1970, 2; M. O. Glenn
collector. 2 66, Cornell University, Ithaca, New York, 16 June 1957, D. R. Davis
collector. ¢, McLean Bogs Reserve, Tompkins Co., New York, 20 July 1957, J. G.
Franclemont collector. ¢, Long Island, New York, no date, Coll. G. P. Engelhardt.
Martha's Vineyard, Mass., no date, ¢; 17 Aug. 1943, 6, F. M. Jones collection. 6, 2,
Oak Station, Alleg. Co., Pa., 4-5 Aug. 1908, Fred Marloff collector. Oneco, Manatee
Co., Florida, 28 March 1954, 2; 23 March 1957, 2; ¢, Archbold Bio. Sta., Highlands
Co., Florida, 27 March 1959, J. G. Franclemont collector. Same location, 28 March
1959, 6; 30 April 1964, 3 66,3 22, R. W. Hodges collector.

North Oaks, Ramsey Co., Minnesota, 7 July 1965, ¢; 16 July 1965, ¢; 20 August
1965, ¢; W. E. Miller collector, in North Central Experiment Station, Saint Paul, Min-
nesota. 6, Livingston Co., Michigan, 22 July 1946, John H. Newman collector, in
Michigan State University. Bay City State Park, Bay Co., Michigan, 17 July 1935, °;
Midland Co., 15 July 1935, ¢, A. Olson & L. K. Gloyd collectors, in University of
Michigan.

Remarks. Nineteen slides, including nineteen genitalia and ten pairs of wings,
have been prepared. These were compared with eighteen slides, including eighteen
genitalia and eight pairs of wings, of Sonia constrictana.

Sonia paraplesiana is extremely close to Sonia constrictana described by Zeller
(1875) from a single male, now in the Museum of Comparative Zoology. This holotype
bears the following labels: 1, Dallas, Texas, Boll; 2, a green label with the hand-written
name Paedisca (?) constrictana; 3, MCZ type No. 14335; 4, Wings and genitalia on
slide V. Adams XII.4.69.

Sonia paraplesiana closely resembles S. constrictana. Zeller’s type (its genitalia and
venation as in Figs. 30, 31, 32) respectively agrees very well with Figs. 18, 20 and 24.

The main characters by which the two species can be differentiated are as follows:

188 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

the outer margin of the forewing of paraplesiana is slightly concave between veins M,
and Cu, and the apex is broadly rounded; in constrictana the outer margin is more
concave between the same veins and the apex is pointed. Sonia constrictana shows
near the apex of the forewing a small ocellus-like spot which on the better marked
specimens is blackish and surrounded with yellowish brown. S. paraplesiana is nor-
mally darkened at the apex of the forewing, but there is no ocellus-like spot there.
Veins Rs and M, of the hind wing are stalked in paraplesiana and the length of the
stalk varies from very short to half as long as the free part of these veins. In constrictana
these veins vary from contiguous to separate and closely parallel for some distance out
of the cell.

The tegumen length of the male genitalia is about equal to the vinculum depth in
constrictana, and greater in paraplesiana; the ventral margin of the valva has a shallow
neck incurvation in paraplesiana, the neck incurvation is deeper in constrictana and
basad of it the margin is bluntly angled. In the female genitalia, the lamella post-
vaginalis of paraplesiana is wider than long and its margins diverge caudad; it is
narrower than long in constrictana and its margins are subparallel. The ventral signum
of paraplesiana is caudad of the dorsal one, they are at about the same level in con-
strictana. The signa of constrictana appear on average, bigger than those of paraple-
siana.

Distribution. Sonia paraplesiana probably inhabits the territory which was indi-
cated by Heinrich for Sonia constrictana: Florida, Texas, North Carolina, Kentucky,
Illinois, Iowa, South Dakota, District of Columbia, Pennsylvania, New Jersey. We can
now add Massachusetts, Michigan, Minnesota, and New York. Sonia constrictana is
more southern: except for one specimen from Putnam Co., Illinois (M. O. Glenn, 1
August 1970, in NMNH), I have seen only specimens from Florida and Texas, where
it is sympatric with paraplesiana and where both species are sometimes taken together.

ACKNOWLEDGMENTS

This paper was made possible through the kind cooperation of
many people: Dr. J. F. G. Clarke, Dr. F. H. Rindge, Dr. R. Silberglied,
Dr. W. E. Miller generously responded to my request for specimens
and types. I am also indebted to Richard L. Brown for comparing
some of my specimens with those in the National Museum and for
several useful comments. I am particularly grateful to Dr. Clarke for
his continued interest and support in this project.

LITERATURE CITED

Brown, R. L. 1973. Phylogenetic systematics: Its application to the genus Epiblema
(Lepidoptera). Unpublished Masters thesis, University of Arkansas. 179 pp.

FORBES, W. T. M. 1923. The Lepidoptera of New York and neighboring states, Cor-
nell University Memoir 68, 729 pp.

HEINRICH, C. 1923. Revision of the North American moths of the subfamily Eucos-
minae of the family Olethreutidae. Bull. U.S. Mus. 123: 298 pp.

KEARFOTT, W. D. 1905. Descriptions of New Species of Tortricid Moths from North
Carolina. Proc. U.S. Nat. Mus. 28: 349-364.

WALKER, F. 1863. List Lepid. Ins. Brit. Mus. Vol. 28.

ZELLER, P. C. 1875. Beitrage zur Kenntniss der nordamerikanischen Nachfalter. Verh.
zool.-bot. Ges. Wien 25: 207-360.

Journal of the Lepidopterists’ Society
33(3), 1979, 189-191

SIX NEW STATE BUTTERFLY RECORDS FROM KENTUCKY!

CHARLES V. COVELL, JR.
Department of Biology, University of Louisville, Louisville, Kentucky 40208

LORAN D. GIBSON
30 Russell Street, Florence, Kentucky 41042

RICHARD A. HENDERSON
2609 Welsford Way, Louisville, Kentucky 40222
AND
MICHAEL L. MCINNIS
8503 Turnside Drive, Louisville, Kentucky 40222

ABSTRACT. Six species of butterflies are added to the Kentucky state list, and
collection data are given. They include Euphyes dion dion (Edwards), E. dukesi (Lind-
sey), Erynnis lucilius (Scudder and Burgess), Calephelis muticum McAlpine, Leptotes
marina (Reakirt), and Lethe portlandia (Fabricius) (subspecies missarkae Heitzman
and dos Passos). A second state record of Poanes viator (Edwards) is also reported,
probably representing subspecies zizaniae Shapiro. The Kentucky butterfly fauna now
numbers 131 species.

The Kentucky Lepidoptera Survey was begun in 1964 with the goal
of characterizing as completely as possible the Lepidoptera fauna of
Kentucky through collection of records and specimens of butterflies
and moths from the Bluegrass State. In a preliminary checklist of
butterflies, Covell (1974) listed 123 species on record as of that year.
Two species have been added more recently to the state list through
published reports: Amblyscirtes belli Freeman (Henderson, 1976)
and Celastrina ebenina Clench (Wagner and Showalter, 1976). While
many exciting discoveries have been made with regard to butterfly
species in Kentucky, notably the discovery of a substantial colony of
Erora laeta (Edwards) in Harlan Co., this paper is confined to the
formal report of six additional butterfly species. These additions bring
the Kentucky list to 131.

Euphyes dion dion (Edwards)

Two males were collected on flowers of Trifolium hybridum L. at
the northern edge of Reelfoot Lake National Wildlife Refuge, Fulton
Co., Kentucky on 29 May 1977, by Gibson and Henderson (one each).
The site was an extensive grass and sedge marsh within a mile of the
Tennessee border. Poanes viator (Edwards) was also taken there on
that date, a second state record of the species; it probably represents
subspecies zizaniae Shapiro. |

1 Contribution No. 194 (New Series) of the Department of Biology, University of Louisville.

190 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Euphyes dukesi (Lindsey)

One male specimen of the Dukes’ Skipper was found by Covell
while examining the collections of the Biology Department, Western
Kentucky University at Bowling Green. The specimen was taken by
student Melinda Johnson in Webster Co., Kentucky on 6 July 1972.
Efforts to locate the student to learn a more precise locality have not
yet been successful. Webster County has much wet meadow habitat
appropriate to dukesi populations. The specimen is now in the Uni-
versity of Louisville collection.

Erynnis lucilius (Scudder and Burgess)

Gerald Straley first reported this skipper from a site 7 mi W of
Irvine, Estill Co., Kentucky, taken 20 April 1974. However, positive
identification has not even now been made. Another possible collec-
tion reported by Straley was taken by him and W. H. Wagner, Jr. 1.2
mi S of Stanton on Route 213, Powell Co., 21 April 1974. Gibson took
4 males and a female in close proximity to the foodplant, Aquilegia
canadensis L. (columbine), along Route 316, Trimble Co., 20 June
1976. Another specimen was taken by Gibson in Bracken Co. amid
extensive columbine on 24 April 1977. Although separation of lucilius
from baptisiae is apparently almost impossible on morphological
grounds (Burns, 1964: 185), we feel that the specimens taken close to
columbine are very likely to be lucilius. Burns (1964) was hesitant to
name specimens from the Appalachian region of Virginia, West Vir-
ginia, and Tennessee as lucilius, feeling that they were “better placed
as baptisiae.” We feel that this ecological evidence justifies including
lucilius on the Kentucky list.

Calephelis muticum McAlpine

A male of the Swamp Metalmark was collected in Otter Creek Park,
Meade Co., 11 September 1976, by Gibson. It was taken on a flower
of tickseed sunflower, Bidens aristosa (Michx.) Britt., in a low, poorly
drained meadow. Covell verified the initial determination on the ba-
sis of forewing snape, coloration, and genitalic features. The specimen
is in the University of Louisville collection (C. V. Covell Male Gen-
italia slide No. 1088). This record extends the known range of muti-
cum to the southeast from central Ohio. In the western extremity of
its range it is known to occur as far south as Arkansas.

Leptotes marina (Reakirt)

One male of the Marine Blue was taken in Louisville, Jefferson
Co., 31 July 1978, by McInnis, and was determined by Covell. The
site was along Interstate Route 71 at Mockingbird Valley Road near

VOLUME 33, NUMBER 3 191

the Ohio River. Crown vetch and other legumes were growing in the
area, but no more specimens were seen on several subsequent visits
to the site. The obvious conclusion is that this individual was a stray.
The specimen is in the University of Louisville Insect collection.

Lethe portlandia (Farbicius)

The number of Lethe species known from Kentucky is now 4 with
the addition of a male and a female of L. portlandia missarkae Heitz-
man and dos Passos on 27 May 1978 by Gibson and Henderson. The
locality was in flat lowland woods with cypress, oak, hickory, and
abundant cane where Route 94 crosses Bayou de Chien, Fulton Co.,
Kentucky. Lethe creola, appalachia, and anthedon have also been
taken in that locality; the creola was abundant on 18 August 1978.
The pair of missarkae was determined by Covell from the original
description. This record marks a new northeastward extension of the
known range of this subspecies, although its presence in Kentucky
had been predicted by Heitzman and dos Passos (1974). The male is
in the Gibson collection; Henderson has the female.

LITERATURE CITED

BuRNS, J. M. 1964. Evolution in skipper butterflies of the Genus Erynnis. Univ. of
Calif. Pubs. in Entomol. 37, 214 p.

COVELL, C. V. JR. 1974. A preliminary checklist of the butterflies of Kentucky. J.
Lepid. Soc. 28: 253-256.

bos Passos, C. F. 1964. A synonymic list of the Nearctic Rhopalocera. Mem. Lepid.
Soc. 1, 145 p.

HEITZMAN, J. R. & C. F. bos Passos. 1974. Lethe portlandia (Fabricius) and L.
anthedon (Clark), sibling species, with descriptions of new subspecies of the for-
mer (Lepidoptera: Satyridae). Trans. Amer. Entomol. Soc. 100: 52-99.

HENDERSON, R. A. 1976. Amblyscirtes belli (Hesperiidae): A new record for Kentucky.
J. Lepid. Soc. 30: 68.

HoweE, W. H. 1975. The butterflies of North America. Doubleday, Garden City, New
York, xiii + 633 p., 97 pls.

WAGNER, W. H. JR. & A. H. SHOWALTER, 1976. Ecological notes on Celastrina eben-
ina (Lycaenidae). J. Lepid. Soc. 30: 310-312.

Journal of the Lepidopterists’ Society
33(3), 1979, 192-196 |

A NEW GHOST MOTH FROM THE SOUTHERN APPALACHIAN
MOUNTAINS (HEPIALIDAE)

DOUGLAS C. FERGUSON

Systematic Entomology Laboratory, U.S. Dept. of Agriculture, U.S. National Museum,
Smithsonian Institution, Washington, D.C. 20560

ABSTRACT. Hepialus sciophanes, a new species from a high elevation habitat in
Jackson Co., North Carolina, is described and illustrated. Its generic affinities are ex-
amined, leading to the conclusion that Hepialus as generally understood is heteroge-
neous. H. sciophanes belongs to the hyperboreus group of northern and western North
America.

Among moths collected in a light trap at nearly 6,000 ft on Water-
rock Knob, Jackson Co., North Carolina, in July 1974 were 19 speci-
mens of a very distinct new hepialid, the first new species of this
family to be described from North America since 1925. It belongs to
the group that includes Hepialus hyperboreus (Moschler), roseicaput
N. & D. (Holland, Pl. 41, Fig. 15, as hyperboreus), pulcher Grt.,
mcglashani Hy. Edw., and mathewi (Hy. Edw.), none of which oc-
curs in the Southern Appalachians. H. hyperboreus, or a species re-
sembling it, has been taken on Mt. Washington, New Hampshire, but
the North Carolina species differs in color, pattern, and male genitalia.
The small, relatively common eastern hepialid, H. gracilis Grt., easily
distinguished by its smaller size and lack of conspicuous white
patches, was the only other hepialid collected on Waterrock Knob.

In an effort to place the new species correctly, I investigated the
European type-species of several genera but reached no satisfactory
conclusion. Members of the hyperboreus group bear a vague resem-
blance to the type-species of Phymatopus Wallengren (hecta L.), hav-
ing comparable although differently developed structures in the male
genitalia. Their relationship to the type-species of Hepialus Fabricius
(humuli L.) and Korscheltellus Borner (lupulinus L.) is less obvious.
Indeed, the male genitalia of these European species and the Amer-
ican ones of the hyperboreus complex are so different that it is hard
to determine whether some of the parts are even homologous. The
new species and others of the hyperboreus group likewise differ
structurally from H. gracilis, H. mustelinus Pack., and from their close
Palearctic counterpart, H. fusconebulosa DeGeer. The latter are
much closer to humuli and lupulinus in the form of all major genitalic
components such as the tegumen, vinculum, valve and juxta, and
agree in the long-stalked condition of R, + R, (almost to apex) in both
wings. Until a much-needed generic revision of the Hepialidae is
available, species of these several groups had best be referred to Hepi-
dlus in the broad sense.

VOLUME 33, NUMBER 3 193

Fics. 1-4. Hepialus sciophanes: 1, holotype male; 2, allotype female; 3, paratype
male, Waterrock Knob, 5,800 ft, Jackson Co., North Carolina, 16 July 1974, D. C. Fer-
guson; 4, paratype male, same data.

Hepialus sciophanes, new species
Figs. 1-6

Description. Venation (Fig. 5) normal for group but differing from that of at least
the most similar eastern species in two respects: R, and R, of both wings stalked for
less than half distance from point of common origin to apex, and R, and R, unstalked
beyond end of cell. In hyperboreus and gracilis R, + R; fork nearer apex, and R, and
R, have a short stalk beyond end of cell.

Males with color and pattern of wings highly variable; basically light brown with
diffuse darker brown lines and shading and a large, elongated white patch in median
area of forewing, inclined obliquely toward tornus. However, over half the specimens
show various degrees of melanism, some heavily suffused with black. Dark markings
of forewing usually obscured, but when visible, consisting of an almost straight sub-
marginal band from near tornus to apex, three costal markings which in higher Lepi-
doptera would be regarded as rudiments of antemedial, medial, and postmedial lines
(in allotype a complete but indistinct medial band present), and diffuse, variable shad-
ing in basal and medial areas that does not resolve itself into definite markings. Pale
markings of forewing consist primarily of the large, white or cream-colored, oblique,
somewhat angulate medial patch, and secondarily of several much smaller whitish
spots as follows: one or more near base, two or three beyond middle of costa, one or
two on inner margin, one or two near mid-zone of submarginal band, one very small
dot near middle of second anal vein, and a subterminal series of five dots. In half of
the specimens the large white patch is reduced or obscured, and in only three are all
white spots present. These white markings are clearly disconnected components of the
pattern seen in more complete form in other species such as H. hyperboreus, mathewi,
or mcglashani, and cannot be reconciled with the very different patterns of H. gracilis
or Sthenopis auratus Grt., the only other southern Appalachian species of comparable
size. Hindwing light brown with thin yellowish costal and outer margins faintly spotted
with brown between vein endings, or darker without yellowish border. Underside of
light specimens brown, with markings of upperside in part showing through faintly

194 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

Fic. 5. Hepialus sciophanes: wing venation of male (paratype).

and with costal and outer margins somewhat yellowish; underside of dark specimens
darker brown, usually without yellowish margins. The allotype, which is the only
known female, is a lighter brown than most of the males, and its pale markings are
suffused with gray. External structure of head, antennae and legs apparently similar
to that of other Hepialidae mentioned.

Male genitalia as illustrated (Fig. 6). Valve well developed, two-lobed. Dorsal end
of tegumen with pair of hairy lobes suggesting socii; a juxtalike plate between bases of
valves, and a medial, basically membranous structure, resembling and probably func-
tioning as an aedeagus, with a distinctive apical sclerite bearing a long, slender spine;
the latter partly encircled by an elaborate, sclerotized structure, in function probably
serving the dual role of gnathos and transtilla, although homologous with neither.
Female genitalia not examined.

Length of forewing. holotype male, 15 mm; paratype males, 15-18 mm; allotype
female, 20 mm.

Types. Holotype, male, Waterrock Knob, 5,800 ft, Jackson Co., North Carolina, 16
July 1974, D. C. Ferguson. Type No. 76,131, U.S. National Museum. Allotype, female,
same data but collected 17 July. Paratypes, 17 males, same locality and collector, 16-
17 July 1974. Paratypes deposited in U.S. National Museum, Canadian National Col-

VOLUME 33, NUMBER 3 195

e

Fic. 6. Hepialus sciophanes: male genitalia (paratype).

lection, British Museum (Natural History), and the collection of Norman B. Tindale,
Palo Alto, California.

Remarks. The type series was collected 500-600 ft S or SW of the Blue Ridge
Parkway, opposite the entrance to the trail leading to the summit of Waterrock Knob,
which is on the other side of the road. The trap was located in a grassy clearing, a small
“bald,” on the mountain slope just outside of a large stand of Fraser fir and yellow
birch that occupies the ridges and summits at that elevation and above. The vegetation
in the immediate vicinity consisted mainly of grasses, ferns, Vaccinium corymbosum
L., Rhododendron and Viburnum species.

Five species of Hepialidae occur in the southern Appalachians: H.
gracilis (H. mustelinus, if indeed a distinct species, may also be
there), H. sciophanes, Sthenopis auratus (easily distinguished by the
extensive gold or silvery markings on the forewing), S. argenteo-
maculatus (Harris), and another large species resembling S. quadri-

196 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

guttatus Grt., known from only one poor specimen from Gatlinburg,
Tennessee. Sthenopis quadriguttatus and argenteomaculatus were
figured by Holland (Pl. 41, Figs. 13, 14). All of these are otherwise
more northern species, or of northern affinity, reaching their southern
limit in the mountains of North Carolina or Tennessee. No Hepialidae
are known from the southeastern Coastal Plain or Piedmont.

LITERATURE CITED

HOLLAND, W. J. 1903 [or later editions]. The Moth Book. 479 pp., 48 pls. Doubleday,
Page & Co., New York.

Journal of the Lepidopterists’ Society
33(3), 1979, 196-197

GENERAL NOTES
A LIST OF LARVAE SUSTAINED ON WHEAT GERM DIET

A wheat germ diet has been formulated for the puss caterpillar (Khalaf 1974, Fla.
Entomol. 57: 377-381). It consists of wheat germ, casein, sugar, salts, inhibitors, linseed
oil, cholesterol, vitamins, antibiotics, and agar. Recently, the same diet was tested and
found suitable for rearing various species of moths. All these, throughout this project,
were collected as larvae from the field after feeding for variable periods on their natural
host plants.

Young larvae of the following species utilized the diet for a period of ten days to a
few weeks and then formed either cocoons or adults: the pyralids, Evergestis rimosalis
(Guenée) and Glyphodes pyloalis (Walker); the notodontid, Schizura unicornis (J. E.
Smith); the noctuid, Spodoptera latifascia (Walker); the arctiid, Diacrisia virginica
(Fabricius); the yponomeutid, Plutella xylostella (L.); the cochlid, Sibine stimulea
(Clemens); the lasiocampid, Malacosoma disstria Hubner; and the liparid, Hemero-
campa leucostigma (J. E. Smith).

Mature larvae of the arctiid, Estigmene acrea (Drury), and the noctuid, Spodoptera
eridania (Cramer) also utilized the diet for about one week or more and then formed
cocoons or adults. More than ten specimens of E. rimosalis, S. latifascia, D. virginica,
S. stimulea and H. leucostigma were reared on the diet. Only a few specimens of the
rest of the species were reared. This diet was slightly modified, mainly by substituting
corn oil for the linseed oil, and was used to rear the puss caterpillar (Khalaf 1975,
Biology of the Puss Caterpillar and its Ichneumonid Parasite. Loyola Univ. Press, New
Orleans, Louisiana. 43 p.). I have used the same modification to raise several other
species of moths: 1) Recently hatched larvae of the noctuid, Spodoptera eridania (Cra-
mer); and liparid, Hemerocampa leucostigma (J. E. Smith) utilized the diet and formed
cocoons or adults. 2) Young larvae of the following species utilized the diet for a few
weeks: the arctiid, Ecpantheria scribonia (Stoll), the notodontid, Schizura unicornis
(J. E. Smith) and the noctuid Spodoptera latifascia (Walker). 3) Mature larvae of the
following species also utilized the diet and formed adults: the lasiocampid, Malaco-
soma disstria Hubner; the aractiids, Diacrisia virginica (Fabricius), Isia isabella (J. E.
Smith), and Hyphantria cunea (Drury); the noctuids, Zale lunata (Drury), Acronicta
arioch Strecker, and Xanthopastis timais (Cramer); and the saturniid, Automeris io
(Fabricius).

In rearing some of the species, e.g., Sibine stimulea, the diet seemed to interfere
with cocoon formation. As the larvae became full grown, I found it was better to plate

VOLUME 33, NUMBER 3 197

only half of the floor of the rearing carton with the diet, leaving the other half clear for
cocoon formation. In other cases, e.g., Spodoptera latifascia and Estigmene acrea, the
mature larvae burrowed into the diet media for pupation. In time, the diet hardened
and trapped the insect. For rearing such species, a special rearing carton must be
prepared for mature caterpillars, half plated with diet and the other half containing soil
for burrowing and pupating.

I am grateful to my students, P. A. Golden, F. J. Mueller, and T. M. Kelly for their
help. I wish to thank R. W. Hodges, E. L. Todd, and D. M. Weisman of the Systematic
Entomology Laboratory, U.S. National Museum, for the identification of the caterpil-
lars. This investigation received support from the Academic Grant Fund of Loyola
University.

KAMEL T. KHALAF, Loyola University, New Orleans, Louisiana 70118.

Journal of the Lepidopterists’ Society
33(3), 1979, 197-198

“MUD PUDDLE CLUBS” IN PURE COLIAS EURYTHEME (PIERIDAE)
IN NORTH CENTRAL CALIFORNIA

Clark (1932, Butterflies of the District of Columbia, pp. 154-155) and Clark & Clark
(1951, Butterflies of Virginia, pp. 109-110) chronicled the invasion and establishment
of the Orange Sulphur, Colias eurytheme Bdv., in the northeastern United States al-
most 50 years ago. They first suggested that “mud puddle” behavior originated in that
species through introgressive hybridization with C. philodice Latr. Puddling by sum-
mer males of C. eurytheme was not observed for about ten years after that species
invaded the Washington, D.C. area. Puddling is still much less common in C. eury-
theme than in C. philodice in the northeast.

One of the few areas in North America where genetically pure C. eurytheme pop-
ulations still exist is the Central Valley of California. The nearest C. philodice are east
of the Sierran divide, north of Mount Shasta, or in the irrigated alfalfa-growing areas
of southeastern California and Arizona (Emmel & Emmel 1973, Butterflies of Southern
California, pp. 18-19). Even so, yellow individuals do turn up rarely in these popula-
tions. They were first noted by Hovanitz (1944, Genetics 29: 1-30) and do not seem to
have increased in frequency in some 35 years, and their origin remains unexplained.

Although C. eurytheme often achieves extremely high densities in the Central Val-
ley, I have been unable to find records of mud puddle aggregations and had never
observed them myself until 26 May 1978. On that date 81 males were counted in four
aggregations in a drying drainage ditch in Rancho Cordova, Sacramento Co., elevation
about 10 m. The aggregations were found between 1455 and 1538 h along 2 km of
ditch; all were in direct sunshine. They consisted of: (i) 10 C. eurytheme, 1 Pieris
rapae L., 3 Everes comyntas Godart; (ii) 23 C. eurytheme, 1 E. comyntas; (iii) 38 C.
eurytheme; (iv) 10 C. eurytheme, 1 P. rapae. All individuals were fresh males. Where
more than one species was involved, each formed a compact group separate from the
others. At another location about 3 km away a single male P. rapae was seen on a
puddle about 1100. Mid-afternoon weather conditions were scattered to broken cirrus
cloud; air temperature 24—27°C, relative humidity 30%; wind SW, ca. 15 km/h.

Nearby annual grassland, occupied by vast stands of a weedy annual Vicia (Legu-
minosae), was the scene of large-scale emergence of C. eurytheme. Virtually all of
several hundred animals seen were fresh males. Many soft-winged individuals could
be found, especially in the morning. A copulating pair, the female teneral, was found
150 m from the ditch at 1525.

198 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Nectar sources were superabundant (blooming Vicia, Brassica, and Centaurea) and
showed no sign of saturation despite the great numbers of butterflies flying. The sample
of male eurytheme I collected from the puddles contained only large, deep orange,
typical “summer” phenotypes. The water content of the vegetation was still high but
beginning to decrease markedly, and the weather had been cooler than normal for a
week. I have no hypothesis to account for the prevalence of puddling at this site on
this date, but it does imply that the capacity to form such aggregations does exist in
pure C. eurytheme.

ARTHUR M. SHAPIRO, Department of Zoology, University of California, Davis, Cal-
ifornia 95616.

Journal of the Lepidopterists’ Society
33(3), 1979, 198-199

LEPIDOPTERA FROM BOLUSES OF NESTLING CATTLE EGRETS
IN EASTERN MISSOURI

The relatively recent immigration and rapid range expansion of the Cattle Egret
(Bubulcus ibis (L.)) in North America (Crosby 1972, Bird-banding 43: 205-212) has
elicited several studies on the feeding behavior and diet of this bird in the United
States (e.g., Hanebrink & Denton 1969, Arkansas Acad. Sci. Proc. 23: 74-79; Fogarty
& Hetrick 1973, Auk 90: 268-380; Jenni 1973, Auk 90: 821-826). The studies that have
been published to date in the U.S.A. mainly provide only the family names of the
arthropod prey species, and their only consensus relative to arthropods is that they are
a major item in the diet of nestling Cattle Egrets. It seems logical to anticipate that
more precise determinations of the arthropod species could help orinthologists des-
ignate certain of the prey species as indicators of portions of the adult Cattle Egret’s
foraging habitat and thereby more accurately interpret the bird’s overall ecology and
behavior.

As part of a preliminary study of the diet of nestling Cattle Egrets, Dr. Jean W.
Graber, Illinois Natural History Survey, collected regurgitated boluses from a heronry
containing ca. 300 active Cattle Egret nests on Billings Island in the Mississippi River,
Scott Co., Missouri, on 30 June and 6 July 1975. Approximately 20 boluses were col-
lected on each sampling date. They were preserved in 70 percent ethyl alcohol and
subsequently examined by me for lepidopterous prey.

The two sets of boluses contained 45 caterpillars and 2 moths representing 9 species
and 4 families (Table 1). The caterpillars in the boluses were in remarkably good
condition thereby facilitating identification of most species. All except two of the cat-
erpillars, Leucania sp. and the single pyralid, are known to be common in field crops
in the midwestern part of the U.S.A. In fact, the species complex suggests that the
caterpillars were captured by adult Cattle Egrets in a legume—grass habitat such as an
alfalfa (Medicago sativa L.) or soybean (Glycine max (L.) Merr.) field. The geometrid
moth, Haematopis grataria Fabricius, frequently may be encountered in short, mixed
herbaceous habitats, e.g., weedy pastures and crop borders (pers. obs.).

Plathypena scabra (Fabricius) primarily feeds on legumes (Pedigo, et al. 1973, J.
Econ. Ent. 66: 665-673) as do the only two species of Colias that occur in eastern
Missouri (see Klots 1960, A Field Guide to the Butterflies of North America East of the
Great Plains. Houghton Mifflin Co., Boston. 349 + xvi p.). The low numbers of Pseu-
daletia unipuncta (Haworth) and Leucania sp., which are both grass-feeding caterpil-
lars (Godfrey 1972, U.S.D.A. Tech. Bull. 1450, 265 pp.) suggests a limited amount of
grass cover in the areas foraged by the adult Cattle Egrets on the dates sampled.

VOLUME 33, NUMBER 3 199

TABLE 1. Lepidoptera from regurgitated boluses of nestling Cattle Egrets (Bubulcus
ibis).

Number Number
of

O
Larvae Adults
Larval 30 7 30. 7
Family Species Foodplants June July June July
Geometridae Haematopis grataria — — — 1 41
Noctuidae Agrotis ipsilon General feeder 8 IW =. =
Caenurgina sp. Grasses, legumes | 1 — —
(crassiuscula or
erechtea)
Leucania sp. Grasses — 1 — —
Plathypena scabra Legumes 1 14 = —
Pseudaletia unipuncta Grasses —- 2—- —
Spodoptera ornithogalli General feeder — § — —
Pieridae Colias sp. (eurytheme Legumes a
or philodice)
Pyralidae Genus ? species ? Unknown 1— — —

Caenurgina species have been associated with both grasses and legumes (Crumb 1956,
U.S.D.A. Tech. Bull. 1135, 356 pp.). Spodoptera orinthogalli (Guenée), a general feed-
er (Crumb 1929, U.S.D.A. Tech. Bull. 88, 179 pp.), is found frequently on soybeans in
the midwestern U.S.A., but not as commonly as Plathypena scabra (pers. obs.). The
polyphagous Agrotis ipsilon (Hufnagel) apparently is associated with moist soil habi-
tats (see Walkden 1950, U.S.D.A. Cir. 849, 52 pp.). Agrotis ipsilon also has been re-
ported as a common lepidopteran species in the stomachs of Cattle Egrets shot near
Cairo and Simbellaween, Egypt (Kirkpatrick 1925, Egypt Ministry Agr. Tech. Serv.
Bull. 56, 28 pp.). This similarity does not suggest that adult Cattle Egrets specifically
forage for the larvae of A. ipsilon, but does indicate that there might be common
denominators in the larval habitat of this noctuid and the foraging habitat of adult
Cattle Egrets that could aid investigations of both species.

GEORGE L. GopFREY, Illinois Natural History Survey, Natural Resources Building,
Urbana, Illinois 61801.

Journal of the Lepidopterists’ Society
33(3), 1979, 199-200

NEW FOODPLANT RECORDS FOR EUPHYDRYAS EDITHA
AND EUPHYDRYAS CHALCEDONA (NYMPHALIDAE)

On 6 June 1978, I located a small colony of Euphydryas editha (Boisduval) near
Frenchman’s Lake in Plumas Co., California. The colony was at about 7,000 ft eleva-
tion, in a dry open meadow transversed by a small semi-permanent creek. The adult -
butterflies (which were well flown and worn), and particularly females, seemed to be
associated with a small yellow plant in the family Scrophulariaceae. A close investi-
gation of this plant, which was later determined to be Castilleja pilosa (Wats.) Rydberg,
revealed two small tents of first-instar (pre-diapause) larvae. The tents contained a

200 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

dozen or so larvae each and they were located near the base of leaves just above the
ground. Euphydryas editha has been known to feed upon other species of Castilleja
(e.g., C. nana Eastw. and C. lapidicola Heller), but not on C. pilosa which is a very
different looking plant to be a “paintbrush.” For a review of the known foodplants of
Euphydryas editha see White and Singer (1:74, J. Lepid. Soc. 28: 103-107).

On 5 June 1978, I was collecting in the Pine Nut Mountains of Douglas Co., Nevada
with David L. Bauer. About 11 mi S of U.S. highway 50, on the Brunswick Canyon-
Sunrise Pass Road, we discovered a large colony of Euphydryas chalcedona (Double-
day) on a dry slope (ca. 5,000-6,000 ft in elevation) in the pinyon-juniper zone. This
is the first time that a colony of E. chalcedona has been located east of the Carson
Valley (although Bauer noted that he had taken a few individual specimens in the Pine
Nuts previously). The butterflies appear to be assignable to subspecies macglashanii
(Rivers), but the usual macglashanii foodplants (Penstemon brevifloris Lindl. and Pen-
stemon lemmonii A. Gray) could not be located in the vicinity; nor could we find any
other known foodplant of E. chalcedona. Adults of both sexes were avidly nectaring
at wild onion (Allium sp.) and we noticed that females were paying quite a bit of
attention to a small Orthocarpus sp. with long filamentous leaves. I caged two females
with several sprigs of the Orthocarpus plant. These females subsequently oviposited
on the plants—demonstrating the probability that this is indeed their foodplant. Pre-
viously recorded foodplants for E. chalcedona have included a number of species of
Scrophulariaceae, including Penstemon, Castilleja, Mimulus, Diplacus and Scrophu-
laria but not Orthocarpus. Thus this is the only report of E. chalcedona making use
of an annual for oviposition. (Is it possible that competition for food between this
species and E. editha, which frequently feeds on annual scrophs, has occurred in other
areas?)

I am indebted to Robert Gustafson of the Los Angeles County Museum of Natural
History for identification of the two foodplants. Because the specimens of the Ortho-
carpus sp. were not in flower, they could be identified only to genus and not to species.

JOHN H. MASTERS, 25711 North Vista Fairways Drive, Valencia, California 91355.

Journal of the Lepidopterists’ Society
33(3), 1979, 200-201

NYMPHALIS MILBERTI (NYMPHALIDAE) NEAR
SEA LEVEL IN CALIFORNIA

Nymphalis milberti Latreille is regarded as a rarity on the Pacific Coast and is usually
recorded at high elevations. Shapiro (1974, J. Res. Lepid. 13: 157-161) pointed out that
it is occasionally taken below 300 m in northern and central California and that such
occurrences seem to involve only overwintered females. It was suggested that N. mil-
berti overwinters at low elevations and breeds there in April, the resulting offspring
dispersing upslope. Because of the low numbers, such movements would be very
difficult to detect. N. milberti, unlike N. californica Bdv., is not considered a migratory
species. Its suggested movements, however, parallel those proposed by Shapiro (1975,
J. Res. Lepid. 14: 93-97) for N. californica. In the northeast N. j-album Bdv. & LeC.
shows a seasonal pattern of occurrence suggesting the same phenomenon (fresh adults
at high elevations in July; overwintered ones at low elevations November—April; Sha-
piro 1974, Search (Agriculture) 4(3); 12).

On 2 April 1978 Mr. Noel LaDue took a worn female N. milberti at Rancho Cordova,
Sacramento Co., California (about 20 m). On 26 May 1978 I took two fresh male N.
milberti on vetch flowers about 1.5 km from the site of LaDue’s capture. Both native
and introduced stinging nettles (potential host plants) occur in the vicinity in riparian
forest. The implication that breeding took place is clear and is bolstered by a report

VOLUME 33, NUMBER 3 201

(J. Brock, in litt.) of early spring breeding near Bakersfield in the San Joaquin Valley
(in another year). It is worth noting that populations of N. milberti were unusually
high in the Sierra Nevada (Nevada, Sierra counties) and the Trinity Alps (Trinity Co.)
in late 1977. On 16 August 1977 several dozen fresh individuals were observed on
Monardella flowers on the south slope of Mount Shasta, 1,425 m. If enough low-ele-
vation records can be accumulated it may be possible to demonstrate regular altitudinal
dispersal even in a species so rare that tagging is unlikely to bring significant results.

ARTHUR M. SHAPIRO, Department of Zoology, University of California, Davis, Cal-
ifornia 95616.

Journal of the Lepidopterists’ Society
33(3), 1979, 201-203

HISTORIS ODIUS (NYMPHALIDAE) SUCKING ON COCOA SEEDS
(STERCULIACEAE) IN NORTHEASTERN COSTA RICA

Although many genera of the subfamily Nymphalinae in the American tropics suck
juices from dung, rotting fruit, and sap flows on trees (e.g., Seitz 1924, Macrolepidoptera
of the World, Vol. 5, American Rhopalocera, A. Kernan Verlag, Stuttgart, 615 pp.; Gil-
bert 1972, Proc. Natl. Acad. Sci. USA, 69: 1403-1407), among the best known for such
behavior are Historis and Prepona. Historis odius Fabricus is a large, robust, and swift-
moving species widespread throughout the West Indies, Mexico, and Central and South
America (Seitz, op. cit.). Although a familiar species in forest light gaps and borders in
tropical rain forest regions generally below 600 m elevation (pers. obs.), H. odius adult
food records are scarce. From the available literature, one must assume that they feed
on rotting fruit, fermenting sap, and dung. While this is very likely the case, I wish to
report H. odius sucking on the drying seeds of the well known commercially-cultivated,
tropical cash crop, Theobroma cacao Linnaeus (Sterculiaceae) commonly known as
“eaecao or cocoa.”

The farm complex Compania Agricola Huntro S.A. (CAHSA) includes extensive
plantings of cacao. During the latter part of the wet season each year, large quantities
of seeds are extracted from pods and placed on drying tables. Before being shipped,
the seeds must be dried. In sunny weather this process usually takes 2-3 days, with
4-7 h exposure each day. When extracted from the pods and placed on the drying
tables, the 30-40 mm long ovoid seeds are individually encased in a white pulp which
is known to be sweet, highly aromatic, and palatable to mammals, which act as dispersal
agents (Cuatrecasas 1964, Contribut. U.S. Nat. Herbarium, 35: 379-614), even though
the seeds are scentless and tasteless to humans. These properties of the pulp apparently
attract small mammals which remove the seeds, suck the pulp, and disperse the seeds
(ibid.). Little is known about invertebrates being attracted to the seeds and pulp. Be-
tween 30 July and 2 August 1978, I had the opportunity to observe H. odius and other
insects visiting the freshly extracted and drying seeds (with pulp intact) at “Finca La
Tirimbina,” a part of the CAHSA complex near La Virgen (220 m elev.), Heredia
Province, Costa Rica. Although the weather is generally rainy and overcast at this time
of the year, the three days of observation were clear and sunny.

At 1100 h 30 July, I noticed three individuals of H. odius sucking on the sticky, moist
surfaces of the drying seeds; all of the butterflies were on the same drying table (about
4 x 5 m) and each had its proboscis wedged down between the seeds (Fig. 1). This
table was shaded under a roof, and other tables in direct sunlight and containing seeds
which had been drying for longer periods had attracted no butterflies or other insects.
Other insects seen on the shaded table included one freshly eclosed Hamadryas februa

bo
(=)
bo

JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

*

ll. a

Fic. 1. Historis odius sucking on the moist pulp of cacao seeds on a drying table
at “Finca La Tirimbina,” La Virgen, Heredia Province, Costa Rica (30 July 1978,
1130 h).

(Nymphalinae) and many Trigona bees (entirely black species). Intermittently through-
out this day and the next one, I noticed that the H. odius adults were present, usually
1-3 at any one time. One adult was very tattered while the other two appeared freshly
eclosed. The latter were extremely wary and would take flight at the slightest distur-
bance, while the tattered individual was more sedentary and could be photographed.
Typically when disturbed, an adult would fly off and perch on a nearby fence or roof
top, only to return to the seeds within 5-20 min. Although all three adults perched on
the seeds within a meter of each other, there were no interactions (aggressive or oth-
erwise) among them. Because of its distinctive and easily recognizable wind damage,
the tattered butterfly seen over a two-day period was assumed to be the same individ-
ual; possibly the same is true for the two fresh adults. By the third day, the seeds were
very dry and the butterflies (and the other insects) ceased to visit them. On the first
day of observation, the seeds were placed on the table by 0800 and had been drying
only a few hours when the butterflies first appeared. During the first day, one fresh
male of Morpho peleides fluttered around the seeds. As my observation periods were
inconsistent, it is not known how many other butterflies exhibited an interest in the
seeds. It was clear that H. odius was feeding on the moist pulp around the seeds. The
drying seeds were highly aromatic to me and to others in the area, presumably the
result of a fermentation process brought on by the drying process.

Owing to the relative inaccessibility of these seeds under natural conditions (encased
in a tough pod or pulp quickly eaten by vertebrates), the pulp is probably not a major
food source. Historis odius and other nymphalines exploit a broad range of rotting
organic substrates in lowland tropical rain forests of Central America. While these
observations occurred during the wet season, H. odius is active throughout the year in
this region since two fresh adults were captured during February and April (1970) at
nearby “Finca La Selva.” Different tribes within the Nymphalinae contain genera that
feed on rotting fruit and tree sap (Howe 1975, The Butterflies of North America, New

VOLUME 33, NUMBER 3 203

York: Doubleday & Co., 633 pp.) and these insects obtain nitrogenous compounds and
other substances that may increase various parameters of reproductive effort or adult
longevity (Gilbert 1972, op. cit.). The timing of visits by the butterflies to the drying
seeds is a behavioral response to products of decay in the pulp. This is a time when
the seeds are highly aromatic but not in the sense traditionally maintained since no
vertebrates appeared at them on the drying tables.

ALLEN M. YOUNG, Invertebrate Division, Milwaukee Public Museum, Milwaukee,
Wisconsin 53233.

Journal of the Lepidopterists’ Society
33(3), 1979, 203-204

NEW OVIPOSITION PLANT FOR EUPHYDRYAS PHAETON
(NYMPHALIDAE)

Larvae of the checkerspot Euphydryas phaeton Drury feed on several species of
Scrophulariaceae and a few other species such as Plantago sp. (Tietz 1972, An index
to the described life histories, early stages and hosts of the Macrolepidoptera of the
continental United States and Canada, vol. 1, Allyn, Sarasota, Florida). However, fe-
male E. phaeton have only been reported depositing eggs on turtlehead (Chelone sp.:
Scrophulariaceae) (Edwards 1884, Butterflies of North America, vol. 2, Houghton Mif
flin, Boston; Tietz, op. cit.). Here I report oviposition by E. phaeton on English plantain
(Plantago lanceolata L.).

I observed four females depositing clusters of eggs on English plantain in an old
field in Manlius, New York on 12-13 July 1978. Twenty-nine egg clusters were col-
lected on plantain: six occurred on two large plants which were touching at the bases,
two were found on the same plant, two others occurred on the same leaf, and the rest
were found singly on plants. Nine of these were on the top side of the leaves rather
than on the under surface. The mean number of eggs in these clusters was 278.7 (range
115 to 516) which was not significantly different from clusters deposited on turtlehead
(C. glabra L.), a larger and broader-leaved plant than plantain (t = 0.00045, P > 0.50,
df = 58) (Stamp, unpubl. data). I spent a total of 6 afternoon hours following females
(n = 7) for periods of 20 to 105 min. All of these females exhibited plant search behavior
for oviposition sites (going quickly from plant to plant), but only two females performed
leaf search behavior (searching plant and touching leaf with tip of abdomen). This
second behavior occurred only on plantain and the first time these females encountered
it during the observation period. The females in this field spent much more time
exhibiting plant search behavior than females observed in areas with turtlehead (Stamp,
unpubl. data). This probably reflects some major differences between the two host
plants. Turtlehead grows 2-4 ft in height, may occur in large, dense patches (diameter
several ft across) and leaves of the plants frequently are touching. In contrast, plantain
is a small plant (height of leaves less than 1 ft), is not common in this field, and occurs
in small patches with plants generally not touching each other. A second population
of E. phaeton in a bog near McLean, New York was also using English plantain for
oviposition sites. Neither of these areas had turtlehead.

The field in Manlius supported a large number of E. phaeton. Using mark-and-re-
capture methods and Bailey’s modification of the Lincoln index (Ehrlich and Davidson
1960, J. Lepid. Soc. 14: 227-229: Poole 1974, An Introduction to Quantitative Ecology,
McGraw-Hill, New York), I estimated 292 adults during this period (peak of flight
season). This was probably only a third of the colony at that time as two adjacent areas
also had E. phaeton and P. lanceolata. One of these areas was a first-year old field in

204 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

which plantain was larger and more common than in the other areas, but it had fewer
adult E. phaeton and only one egg cluster was found there.

It is not surprising that E. phaeton uses P. lanceolata as Euphydryas species in the
western United States use Plantago species for oviposition sites. But it does raise an
interesting question: why aren't there more E. phaeton, especially in old fields, if they
will deposit eggs on such a common plant as English plantain?

I am especially grateful to John Kemper for assisting with the field work. I thank
Edward Jennejohn for showing us the Manlius and McLean populations. This research
was supported by funds from the Univ. of Maryland Chapter of Sigma Xi.

NANCY E. STAMP, Department of Zoology, University of Maryland, College Park,
Maryland 20742.

Journal of the Lepidopterists’ Society
33(3), 1979, 204

PRIOR NAME FOR A PETROVA PINE MOTH (TORTRICIDAE)

Not long after describing Petrova khasiensis (Olethreutinae), a moth developing on
pine in northeastern India, I chanced to see two specimens similar to it at the U.S.
National Museum of Natural History identified as Eucosma argyrocyma Meyrick. Sub-
sequent comparison of pertinent specimens, including male genitalia, confirmed that
the former name is indeed a synonym of the latter. Moreover, both names were based
on specimens from the same locality, the town of Shillong. The findings are summa-
rized below.

Petrova argyrocyma Meyrick, new combination

Eucosma argyrocyma Meyrick 1921, Exot. Microlepidop. 2: 447; Clarke, 1958, Cat.
Type Spec. Microlepidop. Brit. Mus. (Nat. Hist.) Descr. Edward Meyrick 3: 347.
Petrova khasiensis Miller 1977, J. Lepid. Soc. 31: 135, New Synonymy.

Specimens compared included one of the above Petrova argyrocyma (Shillong ...,
5,000 ft 5.28 TBF, Comp. with type... Det.J. F.G.C...., 3 genit. slide MAM 1115781)
and the P. khasiensis 3 holotype. Wing and genitalia illustrations of authentically
determined representatives may be seen in the works by Clarke and Miller cited above.

WILLIAM E. MILLER, North Central Forest Experiment Station, USDA Forest Ser-
vice, 1992 Folwell Avenue, St. Paul, Minnesota 55108.

Journal of the Lepidopterists’ Society
33(3), 1979, 204-205

ABERRANT SATYRIUM A. ACADICA (LYCAENIDAE)

On 4 July 1978 a recently emerged aberrant male Satyrium acadica acadica (Ed-
wards) was captured at Elgin, Kane Co., Illinois. This species is fairly common in the
Chicagoland region wherever the hostplant willow (Salix) grows. The specimen was
collected along a small creek within the city limits of Elgin while it rested on a leaf of
a willow sapling.

VOLUME 33, NUMBER 3 205

Fics. 1-4. Specimens of Satyrium a. acadica from Illinois. 1 and 2, aberrant spec-
imen, dorsal and ventral views, respectively; 3 and 4, same, for a normal specimen,
showing the typical markings of this species.

Figs. 1-4 show the dorsal and ventral views of both the aberrant specimen and a
normal specimen. The most striking difference between them is that in the aberrant
specimen the normal ventral postmedian rows of spots and submarginal “V” markings
are replaced by large rectangular bars extending from the submarginal area into the
postmedian area on both primaries and secondaries. I wish to express my thanks and
appreciation to Dr. Clifford D. Ferris, University of Wyoming, Laramie for making the
photographs of the specimens used in this article.

IRWIN LEEUW, 1219 Crystal Lake Road, Cary, Illinois 60013.

Journal of the Lepidopterists’ Society
33(3), 1979, 206-207

BOOK REVIEW

GozMANY, L. 1978. LECITHOCERIDAE. In Amsel, Gregor & Reisser, Microlepidoptera
Palaearctica, v. 5: text, 306 pp., 168 text figs.; plates, 122 (unnumbered) pp., 15 full
color plates, 93 black and white. Vienna. Price of two bound, gold imprinted parts,
DM540,—(approximately $290.00 U.S.); subscription price, DM450,—.

The new volume of this monumental series fills the reader with admiration: one
holds his breath an instant and wonders that such a beautiful book can be produced in
this day and age.

The series “Microlepidoptera Palaearctica” is so well known by now that my state-
ment that the fifth volume has been written in the same spirit and style, and that it has
been produced at the same high standards as the previous four, characterizes its qual-
ities adequately. So much well-merited praise already has been given to the series,
that it is difficult to add more, without being repetitious. Still, the present volume earns
a special qualification: among those families of Lepidoptera already treated, the Le-
cithoceridae are, without doubt, the least known—and as a separate family, they are
hardly known at all! Amazingly, Dr. Gozmany’s critical revision provides access to a
new group of Lepidoptera in the Palearctic fauna. Therefore, the scientific value of the
fifth volume is considerable.

In the Introduction, an historical survey of the Lecithoceridae is presented. This
group was separated as late as 1947 by Le Marchand as a Palearctic subfamily of the
Gelechiidae, and in 1955 it was described again by Clarke as a separate family (the
Timyridae), which included numerous species of the Oriental and Ethiopian tropics.
Now for the first time the limits of the group are defined and a generic key to the Old
World fauna is given as a basis for its total revision. (The family does not occur in the
New World.) A list of all of the literature on the Palearctic species is compiled and
each species has its own list of citations. Dr. Gozmany was able to study and dissect
all of the type specimens except a very few which could not be found. The limits of
the geographic region have been extended so that several neighboring countries are
included in it [e.g., Nepal, Assam, North Burma and even Formosa (Taiwan)], in order
to incorporate potential future intruding species. This has been done in view of our
still inadequate knowledge of the family. Hardly anything is now known concerning
the early stages and ecology of these insects, except in the case of several common
European species. The Lecithoceridae seem to be chiefly detritophagous, but they also
consume withering and dead plant tissue. Adults are attracted to light. The family is
now divided into two subfamilies, with a total of 41 genera and 168 species. Of these,
15 genera and 67 species are new. Each is illustrated in full color with black-and-
white figures of the genitalia of both sexes. In the second part there are 15 plates of
magnificent watercolors by Dr. Gregor, and additional sketches and genitalia drawings
by the author.

An important novelty is that the description of every species is followed by a special
paragraph, summing up the differences between it and other allied species. Species
discrimination is therefore made possible through keys, diagnoses, color and black-
and-white figures, as well as by these handy summaries—methods worth following in
the future.

With regard to benevolent critical remarks, I have just one: obviously the diagnoses
of the species are meant to be more or less complementary to the color illustrations
(or the other way around); together they are excellent, but separately the diagnoses are
too concise. Perhaps this is a question of taste, but I would have preferred a diagnosis
complete in every detail, as a taxonomic documentation for the identification of each
species.

Entomologists have a strong personal attraction to certain groups of insects. This is
one thing which makes our science so very fascinating! I must confess that I enjoy such
a feeling of involvement with the Lecithoceridae, as well as with certain other groups.
I have been highly privileged to encounter these elegant insects in numbers in south-
ern Asia. This is one more reason that I must congratulate the author, artist, and editor

VOLUME 33, NUMBER 3 207

for their achievements. I welcome the present revision warmly and recommend its
subject to a wide circle of my colleagues interested in Lepidoptera.

Unusual additions to this fifth volume are the sympathetic obituary of the third Editor
of the series, the late Hans Reisser, and a fascinating review of the origin of the ten-
year-old series “Microlepidoptera Palaearctica,’ by its initiator and Editor-in-Chief,
Dr. H. G. Amsel.

A. DIAKONOFF. Rijsmuseum Van Naturlijke Historie Raomsteeg 2, Postbut 9517,
2300 RA Leiden, Nederland.

Journal of the Lepidopterists’ Society
33(3), 1979, 207-208

BOOK REVIEW

EssAI DE CLASSIFICATION DES LEPIDOPTERES PRODUCTEURS DE Solk. Original fascicles
published 1897-1934 in Compte rendu des Travaux du Laboratoire d Etudes de la
Soie, Lyon. Facsimile reprints now available, published 1976-1978 by Sciences Nat,
2 rue André Mellenne, Venette, 60200 Compiegne, France. Price different for each
fascicle, but varying from 42 FF to 99 FF each. Presently only available through
Sciences Nat.

This old classical series on Saturniidae (= Attacidae) and related moths has been
quite rare and unavailable to workers. The series is particularly useful to taxonomists
and of special interest to amateur students of Saturniidae. I own an original copy
of fasc. 2 and can thus see that these reprints are accurate reproductions of the
originals, except for size: originals measure ca. 19 X 27 cm and reprints are ca.
15 X 22 em. It is now possible for both libraries and individual lepidopterists to own
copies of these important works.

The series resulted from the immense interest in these moths from their economic
standpoint as silk producers. Several experts at the Silk Laboratory in Lyon authored
the text, all in French. The figures are not colored, but are line drawings and
some are done from photographs. Almost every species is figured in the adult stage
and a few larvae and cocoons are depicted. Citations to original descriptions and lists
of synonymies precede the text of each taxon. The text mainly describes the imago
and gives the patria. Each fascicle is indexed. The pagination coincides with that
_ of the original separates (extraits) which differs from that in the Compte rendu...
Soie. Below is a synopsis of each fascicle:

Fascicle 1. 1897. By J. Dusuzeau & L. Sonthonnax. Introductory chapter dis-
cussing morphology and early classification of Lepidoptera. Taxonomic group
covered is Saturniidae, Saturniinae, Tribe Attacini: genera Callosamia, Samia
{= Philosamia), Hyalophora, Epiphora, Attacus, Rothschildia, and Archaeoattacus.
Most species figured well. 52 pages.

Fascicle 2. 1899. By L. Sonthonnax. The genus Coscinocera (Tribe Attacini)
which had been omitted from fasc. 1 is covered here. The tailed saturniids are
covered, including the complex of genera related to Saturnia/Eudia; also Agliinae,
Argema, Graellsia, Eudaemonia (= Copiopteryx), Eustera (African). Also included
are Copaxa and Antheraea. Many cocoons and a few larvae are figured. 78 pages.

Fascicle 3. 1901. By L. Sonthonnax. Excepting the Indo-Australian Syntherata
and Neotropical Sagana, all of the species discussed and figured in this fascicle
(such as Nudaurelia and Imbrasia) belong to the large African tribe Bunaeini. 76
pages.

Fascicle 4. 1904. By L. Sonthonnax. A wide range of saturniid groups are

208 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

covered, including the complex of genera related to Saturnia/Eudia; also Agliinae,
Salassinae, more Bunaeini, most Arsenurinae, Polythysana (Hemileucinae), and
Cercophana (Cercophanidae). 86 pages.

Fascicle 5. 1906. By A. Conte. The coverage here is of the large genus
Automeris and its allies, plus a few Nearctic and Mexican Hemileuca. The figures
are quite poor in this fascicle, not having reproduced well. 121 pages.

Fascicle 6. 1908. By A. Conte. The only saturniid genus included is the
African Cyrtogone. Numerous other moths from around the world including many
families, mostly Bombycoidea, are covered. Genera include Borocera, Gonometa,
Pinara, and Taragama. Judging from the lists of synonymies, the literature on many
of these species was scant. 73 pages.

Fascicle 7. 1911. By A. Conte. More non-Saturniidae, as in fasc. 6. Bombyx
mori L. and its relatives, Brahmaeidae, Epia, Ocinara, Synadia, Theophila, Therina,
Endromis, etc. are figured and discussed. 90 pages.

Fascicle 8. 1918. By A. Conte. Supplement to previous fascicles. Text and
figures of over 30 species and subspecies described since publication of the earlier
fascicles. 42 pages.

Fascicle 9. 1931. By E.-L. Bouvier & P. Riel. Contains a catalogue of Saturni-
idae and a list of specimens (and their data) in the collection of the Laboratory of
the Study of Silk. Also a treatise by P. Bonnet on Nephila madagascariensis Vins.,
the great silk-producing spider. Next a chapter on diseases of silkworms by A. Paillot.
Lastly, a chapter on artificial textiles and their chemical and physical properties by
D. Levrat. 141 pages.

Fascicle 10. 1934. By P. Riel. A small second supplement giving text and
figures of 16 Old World Saturniidae. 16 pages.

RICHARD S. PEIGLER, Department of Entomology, Texas A & M University, College
Station, Texas 77843.

Date of Issue (Vol. 33, No. 3): 29 October 1979

EDITORIAL STAFF OF THE JOURNAL
AUSTIN P. PLATT, Editor

Department of Biological Sciences
University of Maryland Baltimore County, 5401 Wilkens Avenue
Catonsville, Maryland 21228 U.S.A.

FRANCES S. CHEW, Managing Editor

Department of Biology
Tufts University
Medford, Massachusetts 02155 USA

oe i, DOUGLAS C. FERGUSON, Associate Editor THEODORE D. SARGENT, Associate Editor

NOTICE TO CONTRIBUTORS

Contributions to the Journal may deal with any aspect of the collection and study of

a Lepidoptera. Contributors should prepare manuscripts according to the following in-

structions.

Abstract: A brief abstract should precede the text of all articles.

Text: Manuscripts should be submitted in triplicate, and must be typewritten,
entirely double-spaced, employing wide margins, on one side only of white, 8% x 11
inch paper. Titles should be explicit and descriptive of the article’s content,including
the family name of the subject, but must be kept as short as possible. The first mention
of a plant or animal in the text should include the full scientific name, with authors
of zoological names. Insect measurements should be given in metric units; times
should be given in terms of the 24-hour clock (e.g. 0930, not 9:30 AM). Underline only
where italics are intended. References to footnotes should be numbered consecutively,
and the footnotes typed on a separate sheet.

Literature Cited: References in the text of articles should be given as, Sheppard

_ (1959) or (Sheppard 1959, 1961la, 1961b) and all must be listed alphabetically under

the heading LITERATURE CITED, in the following format:

SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd. ed. Hutchinson, London.
209 p.

196la. Some contributions to population genetics resulting from the study of
the Lepidoptera. Adv. Genet. 10: 165-216.

In the case of general notes, references should be given in the text as, Sheppard (1961,
Ady. Genet. 10: 165-216) or (Sheppard 1961, Sym. R. Entomol. Soc. London 1: 23-30).

Illustrations: All photographs and drawings should be mounted on stiff, white
backing, arranged in the desired format, allowing (with particular regard to lettering)
for reduction to their final width (usually 4% inches). Illustrations larger than 8% x 11
inches are not acceptable and should be reduced photographically to that size or small-
er. The author's name, figure numbers as cited in the text, and an indication of the
article's title should be printed on the back of each mounted plate. Figures, both line
drawings and halftones (photographs), should be numbered consecutively in Arabic
numerals. The term “plate” should not be employed. Figure legends must be type-
written, double-spaced, on a separate sheet (not attached to the illustrations), headed
EXPLANATION OF FIGURES, with a separate paragraph devoted to each page of illus-
trations.

Tables: Tables should be numbered consecutively in Arabic numerals. Headings
for tables should not be capitalized. Tabular material should be kept to a minimum and
must be typed on separate sheets, and placed following the main text, with the ap-
proximate desired position indicated in the text. Vertical rules should be avoided.

Proofs: The edited manuscript and galley proofs will be mailed to the author for
correction of printer's errors. Excessive author's changes at this time will be charged
to authors at the rate of 75* per line. A purchase order for reprints will accompany the
proofs.

Correspondence: Address all matters relating to the Journal to the editor. Short
manuscripts such as new state records, current events, and notices should be sent to
the editor of the News: Jo Brewer, 257 Common Street, Dedham, Massachusetts 02026
ESA.

PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044 U.S.A.

CONTENTS

THE MALAISE TRAP AS A MEANS OF SAMPLING BUTTERFLY
POPULATIONS IN KENTUCKY. Charles V. Covell, Jr. & Paul
H. Freytag 220220

DIFFERENTIAL GROWTH AMONG LARVAE OF CITHERONIA REGALIS
(SATURNIIDAE) ON THREE GENERA OF FOODPLANTS. C.
Brooke Worth, Thomas F. Williams, Austin P. Platt, & Brian
P. Bradley yo

A DOCUMENTATION OF BIENNIALISM IN BOLORIA POLARIS (NYM-
PHALIDAE). John H. Masters

FOODPLANT OF ALPINE EupPHyDRYAS ANICIA (NYMPHALIDAE).
Raymond R. White 2.

FOURTH ADDITION TO THE SUPPLEMENTAL LIST OF MACRO-
LEPIDOPTERA OF NEW JERSEY. Joseph Muller ___._....__

NEW STATUS FOR EPIBLEMA MINUTIANA (KEARFOTT) AND NEW
SPECIES OF EPIBLEMA HUBNER AND SoniA HEINRICH (TOR-
TRICIDAE). A. Blanchard

S1x NEW STATE BUTTERFLY RECORDS FROM KENTUCKY. Charles
V. Covell, Jr., Loran D. Gibson, Richard A. Henderson &
Michael L. McInnis

A NEW GHOST MOTH FROM THE SOUTHERN APPALACHIAN
MOUNTAINS (HEPIALIDAE). Douglas C. Ferguson

GENERAL NOTES |
Doubly overwintering Citheronia regalis Fabricius (Saturniidae). C. Brooke
Worth
A list of larvae sustained on wheat germ diet. Kamel T. Khalaf ___---__ ee
“Mud puddle clubs” in pure Colias eurytheme (Pieridae) in north central
California. Arthur M. Shapiro

Lepidoptera from boluses of nestling cattle egrets in eastern Missouri. George
L, Godfrey i222) 8 eee rrr
New foodplant records for Euphydryas editha and Euphydryas chalcedona
(Nymphalidae). John H.\Masters. 0.2. J)... 2
Numphalis milberti (Nymphalidae) near sea level in California. Arthur M.
Shapiro 2020050 8 i pe Sail es 1a Oe
Historis odius (Nymphalidae) sucking on cocoa seeds (Sterculiaceae) in
northeastern Costa Rica. Allen\M. Young _..... .
New oviposition plant for Euphydryas phaeton (Nymphalidae). Nancy E.
Stamp

Prior name for a Petrova pine moth (Tortricidae). William E. Miller

Aberrant Satyrium a. acadica (Lycaenidae). Irwin Leeuw

BOOK REVIEWS __

153

162

167

170

174

179

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Sk ee eS Sl eA

q

“

\ a one a

Supplement to

Volume 33 1979 Number 3

JOURNAL

of the

LEPIDOPTERISTS’ SOCIETY

Published quarterly by THE LEPIDOPTERISTS’ SOCIETY

Publié par LA SOCIETE DES LEPIDOPTERISTES
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN
Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS

Fa 29 October 1979

THE LEPIDOPTERISTS’ SOCIETY

EXECUTIVE COUNCIL

T. D. SARGENT, President I. F. B. Common, Immediate Past —
A. M. SHAPIRO, Ist Vice President President ,
ATUHIRO SIBATANI, Vice President JULIAN P. DONAHUE, Secretary

RONALD LEUSCHNER, Treasurer

Members at large:

J. F. EMMEL C. D. FERRIS M. DEANE BOWERS
R. R. GATRELLE J. Y. MILLER E. HODGES
Ay Ps PLEAD M. C. NIELSEN W. D. WINTER

The object of the Lepidopterists’ Society, which was formed in May, 1947 and for-
mally constituted in December, 1950, is “to promote the science of lepidopterology in
all its branches,....to issue a periodical and other publications on Lepidoptera, to
facilitate the exchange of specimens and ideas by both the professional worker and the
amateur in the field; to secure cooperation in all measures’ directed towards these
aims.

Membership in the Society is open to all persons interested in the study of Lepi-
doptera. All members receive the Journal and the News of the Lepidopterists’ Society.
Institutions may subscribe to the Journal but may not become members. Prospective
members should send to the Treasurer full dues for the current year, together with
their full name, address, and special lepidopterological interests. In alternate years a
list of members of the Society is issued, with addresses and special interests. There
are four numbers in each volume of the Journal, scheduled for February, May, August
and November, and six numbers of the News each year.

Active members—annual dues $13.00
Student members—annual dues $10.00
Sustaining members—annual dues $20.00
Life members—single sum $250.00
Institutional subscriptions—annual $18.00

Send remittances, payable to The Lepidopterists’ Society, and address changes to:
Ronald Leuschner, 1900 John St., Manhattan Beach, California 90266 U.S.A.

Back issues of the Journal of the Lepidopterists’ Society, the Commemorative Vol-
ume, and recent issues of the NEWS are available from the Assistant Treasurer. The

Journal is $13 per volume, the Commemorative Volume, $6; and the NEWS, $.25 per
issue.

Order: Mail to Charles V. Covell, Jr., Memoirs Editor, Department of Biology, Uni-
versity of Louisville, Louisville, KY 40208, U.S.A.

The Lepidopterists’ Society is a non-profit, scientific organization. The known office
of publication is 1041 New Hampshire St., Lawrence, Kansas 66044. Second class
postage paid at Lawrence, Kansas, U.S.A. 66044.

Cover illustration: Third instar larva of Limenitis archippus Cramer (Nymphalidae)
preparing to enter winter diapause. The larva is resting on the lip of its hibernaculum
constructed from the basal portion of a chewed tubular willow leaf (Salix babylonica
Linnaeus) covered with silk. In the autumn such larvae begin facultative diapause in
response to decreasing day-length. Original drawing by Mr. George C. Ford, Jr., Graph-
ics Illustrator, Department of Biological Sciences, University of Maryland Baltimore
County, 5401 Wilkens Avenue, Catonsville, Maryland 21228.

JouRNAL OF
Toe LeEPIDOPTERISTS’ SOCIETY

Supplement to Volume 33

Journal of the Lepidopterists’ Society ?)
Supplement to Volume 33

ANNOTATED LIST OF LARVAL FOODPLANT RECORDS
FOR 280 SPECIES OF AUSTRALIAN MOTHS?

NoEL McFARLAND
P.O. Box 56, Northampton, Western Australia 6535?

ABSTRACT

Many new and specific larval foodplant records are listed for 280 spp. of Australian
moths, of which 13 are Microlepidoptera and 267 are Macrolepidoptera; 2 additional
records (1 agaristid, 1 arctiid) are Fijiian. Twenty-two families are represented, as
follows: Agaristidae (8 spp.); Anthelidae (12); Arctiidae, s.]. (11); Carthaeidae (1);
Cochylidae (1); Cossidae (1); Geometridae, s.l. (155 spp. or 55% of the total);
Immidae (1); Lasiocampidae (9); Limacodidae (5); Lymantriidae (4); Noctuidae
(25); Nolidae (10); Notodontidae, s.s. (8); Oecophoridae (3); Pterophoridae (1);
Pyralidae (2); Saturniidae (1); Sphingidae (4); Thaumetopoeidae (11); Xyloryctidae
(4); Zygaenidae (5). Representation of Australian States (encompassing all listed
localities where foodplant records are involved ) is as follows: South Australia (68% );
Western Australia (17%); northeastern Queensland (9%); New South Wales, North-
ern Territory, and western Victoria combined (6%); Tasmania (none). The majority
of S. Aust. records are from the vicinity of Adelaide (Blackwood-Belair district) and
most of the W. Aust. records are from the Geraldton district. Scientific names and
families are given for all plants, with complete author citations and sources of deter-
minations for each. A foodplant index lists most plants by common, generic, and
specific names. All moths are arranged alphabetically in the text, with cross-references
to some generic synonyms. Additional details are given for many spp., usually includ-
ing: (1) code-numbers by which all preserved material can be located in various
named institutions; (2) details on the stages preserved and photographs made for each
life history; (3) dates of adult emergence and of larval occurrence on the plants; (4)
part(s) of the plants eaten; and (5) remarks on distinctive features of larvae or adults
in species that are difficult to separate.

1NOTE: The author regrets that he will not be able to supply reprints of this paper, and that
reprints of McFarland (1972a—1975) are no longer available. This paper is available separately as
a back issue of the Journal of the Lepidopterists’ Society.—Ed.

2 Present address: P.O. Box 1404, Sierra Vista, Arizona 85635.

CONTENTS

INTRODUCTION 2222-2 econnnee ne eee 1
Tue List
Agaristidae 0. se ee 13
Anthelidae ww EEE 14
Arctiidae. Ee 17
Carthaeidae ow err 19
Cochylidae (Phaloniidae) 00.0 SS 20
Cossidae Ee 20
Geometridae —.... Eee 20
Immidae 2.408 a eee AT
Lasiocampidae _......--.. SSS SSS AT
Limacodidae __.....--.-- ee 49
Lymantriidae ‘eee 49
Noctuidae 22228 eer 51
Nolidae 22.1 ee 53
Notodontidae W.... eEe 54
Oecophoridae. _. Ee 56
Pterophoridae .. EEE ee 56
Pyralidae 2. nee 57
Saturniidae te ee 57
Sphingidae ____..._..-- eee 57
Thaumetopoeidae «2a EEE 58
Xyloryctidae (et ee 61
Zygaenidae _ 22 ee ee 62
SUPPLEMENT: Fiji ISLANDS WW. 62
REFERENCES ...--2 2 63
Common NAmMEs oF FoopPLaNTs) 200 eee 66
INDEX TO SCIENTIFIC NAMES OF FOODPLANTS 21... 2 68

ANNOTATED LIST OF LARVAL FOODPLANT RECORDS
FOR 280 SPECIES OF AUSTRALIAN MOTHS

INTRODUCTION

This annotated list summarizes certain information about the adults,
early stages, and foodplants of Australian moths I have reared and studied
over the last 14 years (1 December 1964-30 November 1978). Many of the
species here listed have been reared from eggs which were obtained from
confined females. For most details of the rearing and preservation tech-
niques used, see earlier papers (McFarland, 1964; 1965; 1972a; 1973).
Few (if any ) of these specific foodplant records have ever been published
before, and a high percentage of these life histories were previously un-
known. This work is documented by an abundance of preserved material
(all stages), plus extensive notes and numerous photographs of the living
adults, eggs, larvae, and pupae.

More than half of the foodplant records in this paper involve the family
Geometridae, s.l. (155 spp. or 55% of the total reported here). The
geometrid subfamilies are represented as follows: Ennominae (72 spp.),
Geometrinae (37), Larentiinae (16), Oenochrominae (24), Sterrhinae
(6). Among the other “macro” families, numbers of spp. reported in the
list are: Agaristidae (8), Anthelidae (12), Arctiidae, s.l. (11), Car-
thaeidae (1), Lasiocampidae (9), Limacodidae (5), Lymantriidae (4),
Noctuidae (25), Nolidae (10), Notodontidae, s.s. (8), Saturniidae (1),
Sphingidae (4), Thaumetopoeidae (11), and Zygaenidae (5). Foodplant
records are also included for 13 miscellaneous “micros”: Cochylidae (1),
Cossidae (1), Immidae (1), Oecophoridae (3), Pterophoridae (1),
Pyralidae (2), and Xyloryctidae (4). A number of other foodplant
records had to be omitted due to uncertainty concerning the specific

EDITOR’S NOTE—This issue constitutes a Special Supplement to Volume 33
of the Journal of the Lepidopterists’ Society. Funds to partly cover publication costs
were specially voted by the Executive Council of the Society at the July, 1978 meeting
in Louisville, Kentucky. The present editors wish to acknowledge the able assistance
of Dr. G. L. Godfrey of the Illinois Natural History Survey, Urbana, Illinois, who has
given generously of both his time and expertise in preparing this manuscript for the
publisher. His cooperation is greatly appreciated by both the author and the editors.

2 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

identities of some of the moths, although a few of these have been in-
cluded if the larvae (or the foodplants ) were of particular interest.

This paper was written as foundational to a series of illustrated life
history publications that are intended to follow in the future. However,
this basic information on foodplants and seasonal occurrences is being
reported in view of the man-caused destruction of habitat that is rapidly
spreading through the hills south of Adelaide where most of the listed
species occur. Eventually, all that will remain of the native vegetation
near Adelaide will be the larger trees and shrubs, with most of the smaller
native plants gone; this, in fact, nearly describes the present situation.

Representatives of all South Australian (S. AUST.) specimens discussed
here (material collected between Jan. 1965-May 1970) have been de-
posited in the South Australian Museum of Natural History (Entomology
Department), North Terrace, Adelaide. Other specimens from Western
Australia, and some duplicates from S. Aust., have been deposited in: the
British Museum (Natural History), London; the Australian National In-
sect Collection (A.N.I.C.), Canberra; the Western Australian Depart-
ment of Agriculture (Entomology Section), Jarrah Rd., South Perth.
Most of the Queensland material (1972) has been given to the A.N.LC.
Some of the geometrid larvae are temporarily being retained by the
author for study, but all will eventually be deposited in one or more of
the above named institutions. All of the original photographs (prints
and negatives), and the handwritten notes, plus a complete duplicate
set of the notes (photocopied and stored separately in a bank vault) are
at present being held by the author. Other photocopied sets of the same
notes will eventually be distributed to one or more of the above institu-
tions. A complete index to the collection is also kept up-to-date (in note-
book form), all species being listed in numerical order by family and
code-number; duplicate copies of this index (complete only through
1970) have been deposited at the S. Aust. Natural History Museum
(Adelaide); with D. S. Fletcher at the British Museum (N.H.) (London);
at the Los Angeles County Museum of Natural History (California);
and with T. C. Emmel of the Zoology Department, University of Florida
(Gainesville). This index also covers the North American life history
material in my former U.S. collection (through October 1964), for
which the majority of larval foodplant records were recently published
(McFarland, 1975); this material is in the Los Angeles County Museum
of Natural History.

All specimens in vials have been fixed in K.A.S.A. (see McFarland,
1965) or K.A.A. (3-2-10), and preserved in 80% to 90% ethyl alcohol.
(Ideally, the concentration of alcohol for permanent storage should be
around + 80%; if much lower than 75%, collapse or darkening of speci-

SUPPLEMENT TO VOLUME 33 2

mens will sometimes occur, but if much over 85%, setae often become
too brittle and easily broken. A little glycerine added to the alcohol
may be helpful. )

All reared parasites associated with these moths have been preserved
and deposited in the following three institutions: Diptera mostly in
the A.N.I.C. (Canberra) and the B.M. (N.H.) (London); Hymenoptera
mostly in the W. Aust. Department of Agriculture, Entomology Section
(South Perth). Of the + 40 tachinid spp. reared to date, many have been
determined, at least to genus, by D. H. Colles (Canberra) or R. W.
Crosskey (London), but none of the hymenopterous parasites (+ 60
spp. ) have been identified as yet. All parasites have been code-numbered,
and the details were recorded in two separate series of notes (D and H);
empty puparia and cocoons have always been saved when possible. It is
worth noting that there seems to be a great preponderance of hymenop-
terous over dipterous parasites associated with the geometrids (and
especially the Geometrinae), whereas, in most of the field-collected
larvae representing other families of Lepidoptera, dipterous parasites were
much more frequently encountered. This may have been a coincidence,
of course, but it might reflect the natural situation correctly.

Life History Stages Preserved: At the end of each entry (in paren-
theses ) certain numbers designate exactly which stages of that life history
have actually been preserved and/or photographed. In order to save
considerable space, this information is conveyed simply by use of the
following numbers:

1 = Adults (for photos this usually implies living individuals in their natural
resting positions; sometimes also spread specimens).

2 = Eggs (preserved material can be alcoholic and/or dried unhatched eggs, or
dry empty shells, as described in an earlier paper (McFarland, 1972a: 209-
Pale)

3 = First instar larvae (specifically).

4 — All or some intermediate instars (second through penultimate ).

5 = Last instar (specifically); in notebooks and on all labels this has been ab-
breviated as “L5,” regardless of actual number of instars involved.

6 = Pupae (in alcohol, or empty dry shells inside gelatin capsules, on the pins
beneath reared adults; often both wet and dry are saved).

7 = Cocoons or soil-cells (also occasionally implies larval nests or examples of
feeding-damage to the foodplant, if unusual or distinctive in some respect).
Dry frass samples from last instar larvae of some spp. were also preserved
but have not been mentioned.

8 = Parasites reared and preserved: d= Diptera; h = Hymenoptera; m = mites
or miscellaneous. (Examples: “8dh” would indicate one species of Diptera
and one species of Hymenoptera reared from the same host; “8hhh” would
indicate 3 different hymenopterous parasites from the same host species. )

9 = Notes were made (nearly always including color descriptions based on living
specimens; often details on cryptic coloration, habits, and behavior; also food-
plant and habitat descriptions, etc. ).

4 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

With reference to photographs, the number followed by “c” implies 35
mm color slides (otherwise they are 35 mm black & white). Numerous
habitat photographs were taken between 1966-1970, depicting my main
collecting localities as they were at that time. These are kept in the same
files with the life history photographs. Taxonomists having queries on any
of this material are welcome to write me; I would be able to provide
further details in many cases.

Specimen Code-numbers: A consistent code-numbering system is
vital to this type of work. It is not always possible to quickly identify
reared moths with accuracy (particularly in remote or poorly-known
regions! ), and a percentage usually prove to be new species. The use of
code-numbers guarantees that, as long as specimens or photographs from
my original series exist somewhere, they can always be re-checked by
future workers; these published records will therefore remain essentially
identifiable whether or not subsequent taxonomic revisions cause changes
in the nomenclature, which they certainly must in some cases.

The ennomine geometrid genus Chlenias is a good example: Few of
these relatively similar-looking but variable adults can at present be
easily separated with accuracy, yet the larval differences (in coloration,
maculation, morphology, habits, behavior, and foodplant preferences )
are often consistent and striking, and provide useful taxonomic clues.
The biological and phenological information here reported for Chlenias
spp. took me years to gather and sort out. It would be ridiculous to ex-
clude the unidentified (but obviously different) Chlenias spp. from this
report merely for lack of specific names! Future investigators of the
Chlenias complex will be off to a rapid head start if they have access to
this information and the associated preserved material. The same situa-
tion also applies to the 4 very distinctive but unidentified Pterolocera
larvae (Anthelidae). Code-numbering effectively overcomes the urgency
of immediate naming by providing simple “interim handles” for all taxa
studied. Determinations can follow years later, as convenient (or when
possible ).

All of the rearings, whether from eggs (ex confined 2?) or field-
collected larvae, are code-numbered for separation by family and species.
For example, “G.80” refers only to the oenochromine geometrid (G.),
Phallaria ophiusaria Gn. (80), and not to any other moth. A capitalized
letter (A, B or C, etc.) directly following any code-number implies ma-
terial from another population of the same species. The code-numbers
are enclosed inside all vials containing preserved immatures, and the
identical numbers are handwritten on blue labels attached to the cor-
responding pinned adults. They are also enclosed with all preserved dry
egg shells, frass samples, occasional cocoons, and associated parasites

SUPPLEMENT TO VOLUME 33 5

reared from field-collected larvae. Field notes and related photographs
are also identically code-numbered to cross-reference them to the alco-
holic and/or dry material they represent. In the list that follows, my
code-numbers usually appear in the first line of each entry (after the
determination citation) where applicable; some (few) species were not
code-numbered, usually in those instances where no notes were written.

Dates: The usual or “normal” dates of larval and adult occurrence (in
the localities named) are given, if known with certainty, including peaks
of abundance when these were obvious. All dates applicable to adults
include the entire flight period, from the earliest to the latest recorded
for each species during the study period. All dates given refer only to
the localities named; they are not intended to imply knowledge of flight
times (or larval occurrence ) in any other districts. If the adults came to
light only or primarily after midnight (+ 0130-0400 hrs), a remark to
this effect has been inserted after the months of flight; this applies to
many of the moths studied, even including some winter-emerging species
that fly on cold nights. The ennomine geometrid, Smyriodes aplectaria
(G.140), is an example of the latter.

The months are subdivided simply as follows: “early” = 1st through
fhteamda—tith through 20th; “late°=21st through 3lst. (Finer
subdivisions of the months become meaningless over a period of several
consecutive years in view of the usual local climatic fluctuations from
year to year.) Dates showing months only (minus the year) usually
imply that I have multiple records for that species over several years,
but if the year is also included this usually implies only a single record.

Some (relatively few) of the spring- and summer-emerging moths
have more than one generation per year, whereas the great majority of
autumn- and/or winter-emerging moths are single-brooded, with the
larval stage often extending over two, three, or even more months during
the coldest and wettest part of the year in southern Australia (+ May-
Aug. or Sept.).

Larvae may be presumed diurnal feeders (or feeding both day and
night) unless it is stated otherwise; where included, such information
refers only to final instar larvae. (Some species that feed both day and
night when small become strictly nocturnal feeders in later instars, and
their habits or behavior may also change in other ways.) This distinction,
in the case of strictly nocturnal feeders, can be of great importance when
hunting such larvae in the field. Entirely different searching and collect-
ing techniques are obviously required for larvae inclined to hide by day
under loose bark, well down the stems of their foodplants, or under
debris beneath the plant, often far-removed from their places of nocturnal

feeding.

6 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Abundance-ratings: After “Adults” (in parentheses) a capital letter,
indicating the relative abundance, is given for those species with which
I was more familiar. In the case of Blackwood, this information is based
on numbers of adults attracted to two ultraviolet lights (“black light’—
G.E. F15T8.BL), situated in the same positions during nearly six con-
secutive years of resident collecting in the same suburban garden,
between January 1965 and September 1970. All hours from dusk to dawn
were regularly sampled, during all seasons of the year, throughout this
period.

These abundance-ratings are expressed as follows: (A+) = extremely
abundant; (A) = abundant, (B+) = of moderate abundance but tend-
ing toward (A); (B) =of moderate abundance, not really common but
by no means rare; (B—) = of moderate abundance but tending toward
(C); (C) =rare, with + 6 or fewer individuals seen in most years; (D)
= very rare, with only 2 or fewer individuals seen in most years (some
years none ). These abundance-ratings refer only to the occurrence of adult
moths (at uv. lights), in the specific localities named, and they also
attempt to reflect the “average” or overall relative abundance of cyclic
species during the entire study period. (As in all localities, some South
Australian species fluctuate considerably, whereas others appear to
remain relatively constant in their numbers from year to year. )

The foothills S. of Adelaide are in the path of rapidly expanding de-
struction from mushrooming suburbia (euphemistically known as “de-
velopment’). It can be predicted that changes undoubtedly will take
place in the relative abundance of some of the moths named in the pages
that follow (particularly those restricted to feeding on genera other than
the dominant Eucalyptus or Acacia), as the few remaining vestiges of
“original” habitat disappear around Blackwood, Belair, and elsewhere
in the Mt. Lofty Range. Some of these moths, whose foodplants are
already scarce, may even become extinct in the Blackwood district, or will
inevitably become less abundant than they were in former years. It is
therefore important to keep in mind that most of the observations here
reported were made between 1965-1970.

Localities: Most of these notes were recorded in the Mount Lofty
Range, at 2 Gulfview Rd., Blackwood (elevation 800-900 ft.), a hilly
suburban area about 7 road-miles south of Adelaide, South Australia.
The Blackwood observations were centered almost entirely in and around
the Gulfview Rd. — Hannaford Rd. area. Periodic visits (in all seasons )
were also made to a small but rich and varied remnant patch of nearly
undisturbed native flora inside the northwest section of Belair National
Park, about 2 mi. northeast of Blackwood and between one half to one
mile east of Belair Railway Station, just south of Sheoak Rd.; this habitat

SUPPLEMENT TO VOLUME 33 Th

differed considerably from the Blackwood locality in having a far better
representation of the heath or scrub elements (smaller evergreen sclero-
phyll bushes) of the original Mt. Lofty Range flora.

Western Australian records given as “Drummond Cove, + seven mi. N
of Geraldton” are based on four years of collecting (uv. lights and hunting
for larvae, etc.) at Lot 68, Drummond Cove, or within a short distance
south of Lot 68; this is a coastal sandhill habitat dominated by Acacia
ligulata.

A number of other moth foodplant records are also included from
tropical N. Queensland, the far southwest of Western Australia, and a
few miscellaneous other Australian localities; also, a supplement giving
two foodplant records from the Fiji Islands follows the family Zygaenidae.

All localities named are the exact source-localities of the specimens
(either of the original females from which eggs were obtained or of the
field-collected larvae), regardless of whether or not they were later
transported to some other locality during the process of rearing.

Climate and Seasons: [| interpret the natural seasonal cycle in coastal
South Australia as “beginning” with the earliest soaking rains of autumn,
which is also the cycle in coastal southern California; this is a very
similar Mediterranean climatic pattern, except for the six-month-reverse
in seasons and a somewhat higher (but unreliable) rainfall in the South
Australian spring and summer. The coastal S. Aust. wet season is cool
to cold, followed by a relatively long dry season that is warm to hot with
only occasional precipitation in the form of unpredictable summer
thunderstorms in some years.

In mild-temperate South Australia (Adelaide vicinity), the seasons
can be interpreted roughly as follows: late March to about early May =
autumn (warm to chilly; semi-dry grading to wet); about mid May
through August = winter (mild to cold; mostly wet); September to about
early November = spring (chilly to warm; wet grading to semi-dry);
about mid November to mid March = summer (cool to hot; mostly dry).

Average annual rainfall at Blackwood, S. A., as gleaned from records
kept in the local Post Office (1929-1969), was 26.75”; it had been as low
as 13.71” (1967) and as high as 38.20” (1968), during this 41-year period.
Snow rarely falls, but traces may be seen in the Mt. Lofty summit area
once or twice in most winters. Average monthly rainfall, in inches, has
been as follows (1929-1969): April = 2.44”; May = 3.43”; June = 3.41”;
July = 3.63”; Aug. = 3.21”; Sept. = 2.56”; Oct. = 2.31”; Nov. = 1.62”; Dec.
= 1.26”; Jan. = .89”; Feb. = 1.05”; March = .94”. The seasonal break-
down of this precipitation pattern (rounded off to the nearest inch) turns
out thus: autumn = 4”; winter = 13”; spring = 6”; summer = 4” ( Black-
wood, S. Aust.).

8 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Temperatures: Nocturnal winter lows rarely drop below 34°F.;
diurnal summer highs rarely surpass 102°F. (usually in the 70's, 80’s, and
low 90’s) in the Blackwood district. July is usually the coldest month of
the year (or June-mid August); Jan—Feb. are usually the warmest
months, and can be very hot for brief periods.

Foodplants: All foodplant names listed here have been carefully
checked in the various floras available at the time I was doing this work,
and difficult or variable plants were also submitted to specialists for
determination (see Acknowledgments). Since residence began in W.
Aust. (July 1972), most larval foodplants are now being collected,
pressed, and deposited in the W. Aust. Herbarium (Perth). The family
is named at least once for every foodplant genus, but to save space the
author citations and families of the Blackwood or Belair plants appearing
most frequently in the list are given only once, in the following section
on Vegetation. The system of plant classification employed here (above
the generic level) mostly follows a recent world synopsis of flowering
plants by Thorne (1968); for additional details on this topic, see Mc-
Farland (1970 and 1975: 113). Only standard family name endings
(-aceae) are used. An asterisk (*) preceding the name of any plant
implies that the species is introduced (naturalized and growing wild) in
the locality named, but was not originally native there.

The words “on” or “accepted,” preceding the name of the plant, convey
subtle but important differences: “on” implies that I know (from field
observation) that the plant listed is a natural foodplant of the moth in
the named locality, whereas “accepted” (or “acc.”) implies that the
larvae were reared from eggs in captivity, and that they chose the named
plant from a selection of samples offered by me, but I have no positive
field evidence that they are necessarily to be found on that plant under
natural conditions. However, if the larvae did not thrive upon an offered
plant and successfully produce normal adults, the plant has not been
listed here as “acceptable.” Most of these cases probably do represent the
actual foodplant(s) used in the locality, unless the larvae were reared
in some locality other than that where the original 2 was collected; only
a few cases of this nature are reported here.

The larvae may be presumed leaf-feeders if some other part of the
foodplant is not specifically named. The distinction between young
(new) leaves and mature (old or tough) leaves is often of great im-
portance, particularly in connection with evergreen sclerophyllous plants;
this has been consistently reported if such preferences could be discerned
in the larval feeding habits.

Vegetation: As the majority of records in this paper are from the
Blackwood-Belair district of S. Aust., a brief characterization of the flora,

SUPPLEMENT TO VOLUME 33 fe)

as it was in that locality between 1965 and 1970, seems desirable. The
native plants of the district composed a variable forest-and-scrub mixture
of evergreen sclerophyll trees and shrubs (of many sizes), also including
many grasses, sedges, bulbs, annual herbaceous plants, mosses, and a few
ferns, etc.

Family MYRTACEAE: Eucalyptus odorata Behr. ex Schldl. (pepper-
mint box or mallee box) and E. leucoxylon FvM. (blue or yellow gum
or white ironbark) were the two dominant trees around Blackwood
(especially the former). MIMOSACEAE: Acacia pycnantha Benth.
(golden wattle) was abundant, growing as a large shrub or small tree
under and among the eucalypts. Casuarina stricta Ait. (drooping sheoak,
also a tree) was fairly common in some parts of the district. In the Belair
National Park, S. of Sheoak Rd., Eucalyptus fasciculosa FvM. (pink gum )
was a locally common tree, while at slightly higher elevations nearby,
E. obliqua \Herit. (messmate stringybark) becomes abundant and E.
viminalis Labill. (manna gum or ribbon gum) is not uncommon. At lower
elevations (mainly along watercourses) E. camaldulensis Dehnh. (river
red gum) occurs.

The remainder of the native woody plants were mostly shrubs (many
sizes ), of which the more important species are here listed alphabetically
by family. ASTERACEAE: Olearia ramulosa (Labill.) Benth.; CASU-
ARINACEAE: Casuarina muelleriana Miq. (“oak bush’); DILLE-
NIACEAE: Hibbertia exutiacies Wakefield (syn. acicularis), H. sericea
(R. Br. ex DC.) Benth., and H. stricta (DC.) FvM; EPACRIDACEAE:
Astroloma conostephioides (Sond.) FvM. ex Benth. and A. humifusum
(Cav.) R. Br. (native “cranberry”); FABACEAE: Dillwynia hispida
Lindl., Hardenbergia violacea (Schneev.) Stearn (native “lilac,” a woody
vine), and Pultenaea largiflorens var. latifolia H. B. Williamson;
HALORAGACEAE: Haloragis elata A. Cunn. ex Fenzl and H. hetero-
phylla Brongn. (not woody); LAURACEAE: Cassytha glabella R. Br.
and C. pubescens R. Br. (“dodder’—both are perennial, twining para-
sites of various shrubs ); LORANTHACEAE: Amyema miquelii (Lehm.
ex Miq.) Tiegh. (the common pendulous mistletoe, parasitic only on
Eucalyptus spp. here); MIMOSACEAE: Acacia armata R. Br. ex Ait.
(Kangaroo thorn; common locally, but almost nothing seems to eat it)
and A. myrtifolia (Sm.) Willd. (common understory shrub at slightly
higher elevations nearby); MYRTACEAE (shrubs only): Calytrix
tetragona Labill. and Leptospermum myrsinoides Schldl. (a slender, erect
shrubby “tea tree”); PITTOSPORACEAE: Bursaria spinosa Cav.;
POLYPODIACEAE: 'Pteridium esculentum (Forst. f.) Nakai (brack-

1In Family DENNSTAEDTIACEAE according to Lamp & Collet (1976).

10 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

en)—not woody, but an important perennial where locally common;
PROTEACEAE: Banksia marginata Cay. (“honeysuckle”), Grevillea
lavandulacea Schldl., Hakea rostrata FvM. ex Meisn. (needle bush), and
Isopogon ceratophyllus R. Br.; SANTALACEAE: Exocarpos cupressi-
formis (Labill.) (native “cherry”; a large, shrubby root parasite);
SAPINDACEAE: Dodonaea viscosa Jacq.; XANTHORRHOEACEAE:
Xanthorrhoea semiplana FvM. (Blackboy, grass tree, or yacca; locally
common in the NW corner of Belair National Park).

Of the above-listed native plants, undoubtedly by far the most im-
portant foodplant genus in the Blackwood-Belair locality (throughout
the year) is Eucalyptus, for maximum species of moth larvae feeding
upon it; both new and old leaves are important. A close second in im-
portance (at Blackwood) would be Acacia pycnantha (from June—Oct. ),
but none of the other local acacias. Following these would be a relatively
small number of the other plants, the more important foodplants among
them being Casuarina (both listed species), Hibbertia (only the last two
species listed), Pultenaea largiflorens var. latifolia, Cassytha pubescens,
Amyema, Leptospermum myrsinoides, Hakea rostrata, and Exocarpos.
Of these, one of the most important (from July—Oct.) is Pultenaea
largiflorens, in the Blackwood area; the other plants just named occur
primarily in the NW corner of Belair National Park, or only in very
small and scattered remnant patches closer to Blackwood.

At higher elevations in Belair National Park and elsewhere in the Mt.
Lofty Range, grow numerous other native plants (both woody and
herbaceous), but they are not listed here because these localities are
fairly distant from the area in which most of my collecting was con-
centrated. Some of the moths rated as only “B—” or “C” or “D” in abun-
dance at Blackwood may well be far more common at higher elevations,
or in localities further to the south, where more of the original flora
still grows relatively undisturbed and in more extensive tracts.

Unfortunately, many introduced woody plants (several of them highly
successful and aggressive) are naturalized and extremely abundant in the
Blackwood-Belair district. Notable among these are the following, of
which the first-named is perhaps the worst: Boneseed, Chrysanthemoides
monilifera (L.) T. Norl. (Asteraceae); Canary Is. broom, Genista
(Cytisus) maderensis (Webb & Berth.) Lowe (Fabaceae); furze, Ulex
europaeus L. (Fabaceae); Crataegus sp., Rosa sp., and a blackberry,
Rubus sp. (all Rosaceae); Olea europea L. (olive). Chrysanthemoides
monilifera is steadily advancing into the undisturbed patches of smaller
native plants and is smothering out all smaller species (and even in-
cluding many well-established larger shrubs) as it advances. Unless the
“experts” responsible for the fate of the one excellent small remnant

SUPPLEMENT TO VOLUME 33 1a!

patch of native heath plants still growing a short distance south of
Adelaide (centering about % mi E of Belair Railway Station) suddenly
develop foresight, and become motivated to take the physical action
(i.e., work!) necessary to eradicate the C. monilifera menace from that
area (without harming the native plants ), little will remain there worthy
of preservation in another decade. C. monilifera is, incidentally, one of
the easiest sizeable shrubs to pull up by hand that I have ever en-
countered. I pulled up hundreds in this locality (1966-69)

Of the naturalized weedy herbaceous plants, Arctotheca calendula (L.)
Levyns (capeweed or cape dandelion) (Asteraceae) from S. Africa,
and Echium lycopsis L. (known as “Salvation Jane” in S. Aust. and
“Paterson's Curse” in W. Aust.) (Boraginaceae), are among the most
abundant and conspicuous species growing in disturbed open places
throughout the district. Other conspicuous weeds here are several S.
African Liliaceae (bulbs), Oxalis pes-caprae L. (Soursob), Plantago
lanceolata L., Polygonum ?aviculare L. (wireweed), and various intro-
duced grasses.

Acknowledgments and Determinations: Al] names used in the list
that follows were based on the associated adult moths (those bearing code-
numbers on blue labels). Many of these moths have been matched with
the type specimens. Sources of determinations used here (both ento-
mological and botanical) are abbreviated as follows: BH =B. Hyland
and A. Irvine of the Forestry Regional Research Station, Atherton,
Queensland (N. Qld. plants); IC =I. F. B. Common of the Australian
National Insect Collection, C.S.I.R.O., Canberra, A.C.T., helped with
the majority of moths in this study (all families); MK =the late Mrs.
M. Kenny of the South Australian Museum, Adelaide (S. Aust. plants);
NM =the author (some specimens so designated were also submitted to
specialists for verification); SF = D. S. Fletcher, and his assistant, Mrs.
K. Smiles, of the Entomology Department, British Museum (Natural
History ), London (primarily geometrids and some miscellaneous moths );
Fletcher also reviewed the manuscript in 1973 and offered helpful sug-
gestions. WAH = various taxonomists of the Western Australian Her-
barium, Jarrah Rd., South Perth (since 1968), and P. G. Wilson in particu-
lar (W. Aust. plants ).

I am deeply indebted to D. S. Fletcher and I. F. B. Common for their
frequent and generous assistance, since 1965, with numerous difficult
moth determinations, sometimes involving the preparation of genitalic
slides and thus the expenditure of much time on my behalf; without their
help, publication of these records would have been very difficult. In
the case of my own determinations (NM), they were obtained primarily
from studying material in the extensive South Australian Museum moth

12 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

collection (Adelaide) while I was on the staff there, supplemented by
careful examinations of other major collections in Melbourne, Sydney,
Canberra, Brisbane, and London (BMNH only). My interpretation of
moth families (and superfamilies) follows Common (1970: 752-83).
Plant determinations (other than above-credited) were obtained pri-
marily by careful keying in the floras available for the various regions
at the time of this study (see References). Mr. T. R. Newbery (TN),
of Murray Bridge, S. Aust., kindly provided a number of his foodplant
records, which are included in the list; his companionship in the field,
upon numerous occasions, will always be remembered with pleasure.
Similar pleasant memories are also associated with past field trips in
the company of Mr. K. J. Sandery (St. Agnes, S. Aust.), and the late
Mr. and Mrs. J. O. Wilson of Glenelg (see McFarland, 1974). The
Sandery and Wilson moth collections, while not large, contained many
specimens of interest. The Wilson Collection has gone to the A.N.IL.C.
I would also like to thank my former landlady, Mrs. L. Henley,
for her unfailing tolerance of my strange nocturnal pusuits in her
back garden during my nearly 6 years of residence at 2 Gulfview Rd.;
my wife, Dienie, for assisting at home and afield in various aspects
of this work, for cheerfully sharing her home with hundreds of lepi-
dopterous larvae during the first years of her marriage, and for expertly
typing this manuscript. Appreciation must also be registered for her
patience in coping with a grossly overloaded refrigerator, which was
periodically choked with plastic bags containing larval foodplants!

Abbreviations & Symbols: acc. = accepted; det. = determined by;
fl., fls. = flower(s); fr., frs. = fruit(s); “H.” (before a date) implies
the date of adult emergence from a pupa in captivity; If. = leaf; lvs. =
leaves; nr. = near; “orig. 2?” = the original female from which eggs were
obtained in confinement; + means more-or-less, about, or approximately;
an asterisk (*) before the name of a plant (or moth) indicates that the
species is introduced or not originally native to the locality named. For
convenience and quick reference, all moths are arranged alphabetically
from the family down. Australian states are abbreviated as follows in
the list: QLD. = Queensland; N.S.W. = New South Wales; N. TERR.
= Northern Territory; S$. AUST. = South Australia; TAS. = Tasmania;
VIC. = Victoria; W. AUST. = Western Australia.

Terminology for Adult Resting Positions: The following terms are
used for the three major categories of resting positions commonly seen
in adult Macrolepidoptera: (1) tectiform =the forewings held more-or-
less roof-like over the abdomen, upper surfaces exposed, with the hind-
wings more or less completely hidden beneath (seen in many noctuids
and arctiids, etc.); (2) planniform = all wings more or less flatly ap-

SUPPLEMENT TO VOLUME 33 123

pressed to the substrate, commonly with some degree of hindwing ex-
posure (seen in numerous geometrids ); (3) veliform = the typical butter-
fly position of “total rest,” with all wings held erect over the dorsum, the
upper surfaces tightly closed together, sail-like. The veliform position is
also seen in a few geometrids such as the Australian oenochromine,
Hypographa aristarcha, and the North American genera Fernaldella and
Stamnodes, etc. Xanthorrhoe and Hydriomena spp. also sometimes exhibit
this position temporarily (when just landed), but usually revert to
planniform later (when at “total rest”). Aside from a few geometrids, I
know of no other Australian moths that use the veliform position.

The first term is well-known, but I have never seen the other two used
before, so am proposing them here. There are numerous subtle variations
on these three major themes, which can usually be adequately described
and compared simply by indicating the degree of exposure of the hind-
wing; this can be expressed as a percentage. Any rolling or folding of
wings or margins should also be noted, as well as any tendency to clasp
a branch or twig with the forewings (for anchorage). Examples of the
latter are often encountered in windy climates. Among others, Australian
ennomine geometrids of the genera Capusa, Lophothalaina, and Stathmor-
rhopa have this clasping habit well developed. When at rest, Capusa spp.
also partially fold the forewings fan-like, in a most peculiar and distinctive
fashion, which causes them to appear much narrower distally than they
actually are when the wings are spread.

{bigs Joe

NOTE: All plants not showing family names or author citations in the list below
are discussed under Vegetation (pp. 9-11), where this information is given once
in full for each of these common South Australian species.

AGARISTIDAE

e Agarista agricola (Don.) (det. MM)—As.12. N.QLD., nr. Ravenshoe, at Mill-
stream Falls (J. Wrigley, collector): Larvae (May—June 1972) conspicuous on a vine,
Cissus sp. (probably C. opaca FvM.); captive larvae accepted (as a substitute)
another vine, Cayratia ?clematidea (F. Muell.) Dom.—both VITACEAE (dets.
BH). Adults diurnal. See Common (1966b: 71-color; 1970: 863) for illus. of adult.
(Preserved = 5, 9; photos = 5.)

e Apina callisto Walk. (det. NM)—As.8. S.AUST., Adelaide, suburb of Walker-
ville, just NW of Kingston Tce. at Francis St., in the city parklands (NM, collector ):
Larvae (late July—early Sept.) extremely abundant only within the limits of a small
and restricted “colony,” in a large grassy-weedy field (paddock), feeding primarily
on the S. African annual weed, Cape dandelion, *Arctotheca calendula; a few larvae
seen feeding on two other low-growing weeds, *Malva sp—MALVACEAE, and
*Rumex sp.—POLYGONACEAE; one larva seen feeding on a *grass—POACEAE
(dets. NM). Larvae are diurnal feeders and need sun to elicit vigorous feeding.

14 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Adults (B+) fly mid April-mid May (univoltine) and are strictly diurnal; flight
mostly confined to very restricted areas of the general habitat. See Common (1966b:
121) for photos of larva and adult; McFarland (1970: 350 & 1972b: 227) for egg
photos. (Preserved = 1-6, 8dd, 9; photos = 1, 2, 6.)
@ Comocrus behri (Angas) (det. NM)—As.5 & 5A. (1) S.AUST., Blackwood-
Belair district (NM): Larvae (Jan.—April) conspicuous (but widely-scattered) on
the pendulous mistletoe, Amyema miquelii (det. NM), which is parasitic on many
gum trees, Eucalyptus spp. Adults (B) fly late Nov._March (peak Jan.); diurnal
only. (2) S.AUST., + 30 mi. S of Whyalla, at Murninnie Beach (B. Flounders,
collector); Larvae (April 1969) common on Amyema melaleucae (Lehm. ex Mia. )
Tiegh., which was parasitizing Melaleuca lanceolata Otto—MYRTACEAE (dets.
MK). For a discussion of diurnal hilltopping behavior in W. Australian adults of this
sp., see McFarland (1976). See Common (1966b: 121) or Tillyard (1926: Pl. 36)
for adult photos. (Preserved = 1, 5, 6, 9; photos = 5, 6.)

Cremnophora angasi Walk.—see Noctuidae.
e Hecatesia exultans Walk. (det. IC)—As.11A. W.AUST., 22-24 mi. ESE of Hyden,
at roadside (D. & NM): Larvae (late Oct.) conspicuous, sunning and feeding on the
young (red) growing tendrils of devil’s twine or Australian “dodder,” Cassytha sp.,
which was commonly parasitizing a shrubby Casuarina sp., as well as other plants in
the area (dets. NM). Adults fly in spring and summer; peak of ¢ activity probably
crepuscular; peak of @ activity probably evening and/or nocturnal. Sexual dimor-
phism is apparent. (Preserved = 1, 5, 6, 8d, 9.)
e Hecatesia thyridion Feisthamel (det. SF )—As.9 & 9A. (1) S.AUST., Mt. Lofty
Range, Upper Sturt, nr. Ironbank Rd. (NM, TN, & P. Taverna): Larvae (Feb.—
April) at rest and feeding (diurnally) on young tendrils of Cassytha pubescens and
C. glabella (dets. MK), parasitic twiners on many shrubs in this locality. Adults (B,
very local) fly Jan._March; peak of ¢ activity probably crepuscular, after hot days;
peak of 2 activity probably nocturnal (occasional individuals of both sexes come to
uv. light). There may also be a spring emergence (Oct—Nov. +). Sexual dimor-
phism is apparent. See Common (1966b: 123) for ¢ adult photo; Common (1970:
Pl. 8, color) for g adult of H. fenestrata Bdy. (Preserved = 1, 2, 5, 6, 8d; photos =
2, 5, 5c, 6.) (2) W.AUST., 13 mi. E of Tambellup, at roadside (NM & N. B. Tin-
dale): One larva (21 Nov. 68) on young growth of Cassytha sp., which was para-
sitizing Melaleuca uncinata R. Br. ex Ait—Myrtaceae (det. NM). (Preserved =
D9),
e Periscepta polysticta (Butl.) (det. IC)—As.7. S.AUST., S coast of Kangaroo
Island, 2% mi. S of Mt. Taylor (+ 10 mi. W of Vivonne Bay), on land of G. D.
Seton (NM): Larvae (Dec.—Jan.) locally common on the dwarf shrub, Hibbertia
fasciculata R. Br. ex DC. (det. MK). Adults (B) fly Oct.—Nov., diurnal only. See
Common (1966b: 123) for adult photo. (Preserved = 1, 5, 6, 9; photos = 1.)
@ Phalaenoides glycine Lew. (det. NM)—As.6. (1) S.AUST., Blackwood (NM):
Larvae (Oct.—April) common on cultivated grape vines, *Vitis spp.—VITACEAE;
also several in garden (Nov. 69) feeding avidly on lvs. of an introduced Californian
evening primrose, *Oenothera hookeri T. & G—ONAGRACEAE (dets. NM).
(2) S.AUST., N of Penola (NM & D. Lee): Larvae (Dec. 64) on a small annual
Epilobium sp.—ONAGRACEAE (det. NM), at the edge of a road. (Interestingly,
this same contrasting combination of foodplant families, although involving various
plant genera and species, is also known for some of the North American agaristids and
sphingids.) Adults (B) fly mid Sept.-Mar. (peak Nov.—Jan.), diurnal only. See
Common (1966b: 121) for adult photo; Common (1970: 858) for line drawings of
larva and pupa. (Preserved = 1, 5, 6, 9; photos = 5, 6.)
e Sarbanissa (Seudyra) bostrychonota Tams—see Supplement on Fiji (following
Zygaenidae ).

ANTHELIDAE

© Anthela denticulata Newman (det. NM)—An.1. S.AUST., Adelaide and Black-
wood districts (NM): Larvae (July—Sept.) locally common on various naturalized

SUPPLEMENT TO VOLUME 33 15

and native grasses (esp. “soft” annual spp.) —POACEAE, in weedy pastures, vacant
lots, along roadsides, or in comparable situations. The larvae are diurnal feeders and
seek the sun. Adults (B) fly mid March—April, especially after 2300 hrs; univoltine.
(Preserved = 1-7, 9; photos = 2.)

e Anthela glauerti Turner (det. IC)—An.19. W.AUST., + 125 mi. S of Carnarvon,
at the “Overlander Roadhouse” (D. & NM): Orig. @ (13 July 77) had come to
lights the night before. Readily oviposited in a jar (on a muslin strip). Captive
larvae (reared at Drummond Cove, W.Aust.) readily accepted Acacia ligulata (det.
NM); new lvs. preferred. A second generation mating of this very small anthelid
was obtained in captivity (Sept. 78), and some of these eggs were frozen for perfect
specimens before hatching; 1977 eggs were preserved as empty (hatched) shells.
Mbresenved = 6-9 1, 2, 3-5P, 6, 7?.)

e Anthela ocellata (Walk.) (det. NM)—An.4. S.AUST., Adelaide city suburbs, in
gardens and vacant lots, etc.: Larvae (Nov.—June) seen feeding on various natural-
ized annual grasses, and on mixed lawn grasses—POACEAE. Adults (B) fly spring—
autumn. This sp. appears to be far more abundant in city gardens, parks, and other
cultivated areas than it is in areas where the native flora predominates. See Common
(1966b: 93) for ¢ adult photo. (Preserved = 1-7, 9; photos = 2.)

e Anthela sp. nov. (det. IC); close to repleta Warren (det. SF)—An.7 & 7A. (1)
S.AUST., 5 mi. E of Two Wells, at roadside (NM & TN): Larvae (Aug.—Sept.) on
Cassia nemophila Cunn. ex Vogel—CAESALPINIACEAE (det. MK). (Preserved =
1, 4-6.) (2) S.AUST., Blackwood-Belair district (NM): Larvae (Sept.—Oct.) rest
in full view on young lvs. of golden wattle, Acacia pycnantha (det. NM). Adults (B)
fly late April-July (peak June), often on very cold winter nights when little else
is on the wing. ¢ & @ quite diff. in color (@ usually much paler); univoltine.
(Preserved = 1-7, 9; photos = 1, 2.)

e Anthela sp.; close to (but not) guenei Newman (det. IC)—An.11. S.AUST., at
Kingoonya, approx. 400 mi. NW of Adelaide (M. McFarland, NM, & V. Lill):
Larvae (late March) common on Cassia nemophila vars. and one other Cassia sp.—
CAESALPINIACEAE (dets. NM); these plants were full of luxuriant new growth, in
response to heavy rains several weeks earlier. Adults probably fly between Sept. and
March, whenever rains break the pupal diapause during warm or hot weather (a
desert locality, with variable and unpredictable rainfall). Reared specimens from
this series are also in the K. D. Fairey & V. J. Robinson collections. (Preserved =
1, 5-7, 9.)

e Anthela sp., probably xantharcha Meyr. (det. NM)—An.12 & An.18. S.AUST.,
inland desert, 4% mi. NNW of Coober Pedy, in a small, rocky, dry creek bed
(M. McFarland, NM, & V. Lill): Several conspicuous masses of large opaque, pure
white eggs (23 March 69) were found attached to lvs. in the tops of Eremophila
freelingii FvM.—Myoporaceae (large bushes), almost certainly not one of its
foodplants; a Cassia sp. (the probable foodplant) was growing nearby. Captive
larvae (reared at Blackwood, S.Aust.) readily accepted young lIvs. of Acacia
pycnantha (dets. NM) and grew well. They entirely refused lvs. of a few Myoporum
and Eremophila spp. offered to them, but E. freelingii was not among the samples.
For notes on the early stages of a W. Australian xantharcha population, see Mills
(1954). (Preserved = 1-7, 8h ex egg; photos = 2.)

e Anthela sp. (det. IC)—An.14. (1) N.QLD., + 9 mi. W of Mareeba (J. Wrigley):
Larva (3 June 72) on mature lvs. of a poisonous tree, Cooktown ironwood, Erythro-
phleum chlorostachys (F. Muell.) FABACEAE (det. BH). The resulting ¢ adult
(H. 23 Sept. 72) and its cocoon and pupal shell are in the A.N.I.C. (Canberra).
(Preserved = 1, 4, 6, 7, 9.) (2) N.QLD., 16 mi. SW of Conjuboy Homstead
(+ 2400’ el.), N of Hughenden (D. & NM): Nearly fullgrown larva (21 June 72),
identical to the above, at rest among mature lvs. of E. chlorostachys, + 12 feet up
in the tree. (Preserved = 5.)

e Munychryia periclyta Common & McFarland (1970) and M. senicula Walk. (det.
IC )—“N.”109. Larvae of both are feeders upon the foliage of various Casuarina

16 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

spp. and are described in detail (including life history photos of all stages of M.
senicula) by Common & McFarland (1970). “N.”109 refers only to senicula (in S. A.
Museum). Adults of senicula (Belair district, S.AUST.) fly in spring-summer. See
also Common (1970: 850) for ¢ adult photo. (M. senicula only: Preserved =
1=79: photos: = 1; 274-7.)

e Pterolocera sp., close to amplicornis Walk. (det. NM)—An.6 & 6A. (1) S.AUST.,
Hallett Cove, coastal bluffs + 10 mi. S of Adelaide (D. & NM & K. D. Fairey):
Larvae (June—Aug.) extremely common locally, most often on (or very near) the
ground, concentrated in suitable habitats, sunning and feeding (diurnal only) on
low-growing plants, primarily on tender naturalized (annual) and tough wiry (peren-
nial) native grasses—POACEAE;; also seen feeding (occasionally) on the very tough-
wiry lvs. of Gahnia lanigera (R. Br.) Benth—CYPERACEAE (dets. NM). These
larvae drink (dew or rain) copiously at times, and they also require plenty of fresh
air, as well as sun (periodically); if any of these needs are totally ignored they will
not thrive in captivity. Adults (¢ ¢ only) have a brief but concentrated flight in
autumn (+ April), when they search for the wingless 9 2. Sexual dimorphism is
spectacular. See Common (1966b: 241 or 1970: 850) for ¢ adult photo of ampli-
cornis; McFarland (1970: 349 & 1972b: 223) for egg photos. (Preserved = 1, 2,
4-7, 8ddd, 9: photos = ¢1, 2,5, 6 96,7.) (2) S.AUSTE, Si coasticrKeinganoosls=
at Seal Bay (NM, M. Pate, & G. D. Seton): Larvae (16 Oct. 66) abundant, wander-
ing over the sand in association with larvae of my Ar.34A (Arctiidae), and the even
more abundant Lp.12 (Lymantriidae); seen feeding (diurnal) on various small
herbaceous spring ephemerals, as follows: Crassula sp—CRASSULACEAE; Daucus
glochidiatus (Labill.) Fisch., Mey., & Avé-Lall., and Hydrocotyle sp.—both
APIACEAE; also one eating (perennial) *Poa bulbosa L.—POACEAE (dets. MK).
(Preserved = 1, 5, 6, 8d, 9.)

e Pterolocera sp. (det. IC)—An.2 & 2A. (1) S.AUST., Mt. Lofty Range, in Belair
Nat. Park, just S of Sheoak Rd. (NM): Larvae (July—Oct.) feeding both at night
and diurnally, on many unrelated plants, both woody shrubs and tough, wiry perennial
bunch-grasses. Specific records: Older larvae (in this locality) seem to show a
preference for nocturnal feeding on the tough, needle-like, sclerophyll old lvs. of
Hakea rostrata; one seen feeding (mid Aug.; day) on Casuarina striata Macklin.
(dets. MK). Adults (¢¢ only) have a brief but concentrated flight in autumn
(mid March-April or May), when they search for the wingless ? 9. Sexual dimor-
phism is spectacular. (Preserved = 4, 5, 9.) (2) S.AUST., Kangaroo Is., 2% mi.
S of Mt. Taylor (+ 10 mi. W of Vivonne Bay), on land of G. D. Seton (NM):
Larvae (mid Oct.) abundant, but widely-scattered throughout the native (sclero-
phyll) scrub of this locality; in daytime usually up in various bushes, not on the
ground (unlike An.6 & An.6A); feeding both day and night on the tough, mature
lvs. of their foodplants. Specific feeding records (based on field observations ):
Casuarina striata Macklin.; Hakea Pmuelleriana Black; Daviesia brevifolia Lindl. and
Platylobium obtusangulum Hook.—both FABACEAE; Choretrum spicatum FyM.—
SANTALACEAE (dets. MK). Larvae brought back to Blackwood readily switched
over to Acacia pycnantha as a substitute foodplant. (Sun not imperative for this
species in captivity, but ample ventilation is desirable—also an occasional sprinkling. )
See McFarland (1970: 349 & 1972b: 223) for egg photos. (Preserved = 1-7, 8h, 9;
photos = 91,2) 4.96)

© Pterolocera sp. (det. IC)—An.10. W.AUST., Stirling Range, along roadside nr.
Toolbrunup Peak, + 200 mi. SE of Perth (NM & N. B. Tindale): Small larvae (19
Novy. 68) fairly common on young lvs. of (only) the low shrub, Banksia sphaerocarpa
R. Br. (det. RR). Readily accepted young lvs. of B. marginata as a substitute food-
plant, when taken 1200 mi. E to Blackwood; grew to full size, in excellent condition,
on the latter. (No apparent urgent need for sun in captivity, but ample ventilation
and a light sprinkling every day or two are desirable.) The adult @? is wingless;
sexual dimorphism is spectacular. This sp. shows a kinship with my An.2, in the

SUPPLEMENT TO VOLUME 33 17

general appearance of adults, the egg shape (profile), and the larval behavior, but
it is easily separated and is probably a distinct species.

Mites: I have never seen any lepidopterous larvae more heavily mite-infested

than were some of the younger individuals of this sp. when first collected! (See
also remarks under Carthaeidae.) These vivid red mites were later det. by Dr. R. V.
Southcott (Adelaide) as Charletonia feideri Southcott, 1966 (ref. Aust. J. Zool. 14:
752) of the Erythraeidae, Sbf. Callidosomatinae, Tribe Charletoniini. (Preserved =
1-7, Sm, 9; photos = 2, 5.)
e Pterolocera sp. (det. NM)—An.20. W.AUST., Moresby Range, + 19 mi. NNE
of Geraldton, at Howatharra Hill Reserve, especially in Zones 5 and 10 (D. & NM):
Larvae (Aug.—Oct.) abundant most years, usually seen feeding, sunning or at rest on
grasses or sedges, esp. Lepidosperma. The short middorsal setae form a conspicuous
pinkish-white line in this population; I have not as yet reared or collected any adults.
These larvae are clearly distinct from An.10, but might prove to be a ssp. of An.6 or
An.2. (None preserved at time of writing. )

General remarks on the foregoing Pterolocera spp.: These 4 incomplete determina-
tions were included in the list because of the good differences in larval appearance,
morphology, coloration, behavior, and foodplant preferences; also, the eggs are
easily separated by differences in the coloration, maculation, and measurements
(proportions). The ¢ adult moths, however, are of rather similar appearance, some-
what variable in coloration, and sometimes inclined to overlap in this variability;
the species are very often mixed up in collection series. All four are univoltine.

ARCTITIDAE
(A) SuBFAMILY ARCTIINAE

e Amsacta marginata (Don.) (det. IC)—Ar.46. W.AUST., Northampton (NM):
Larvae (July—Sept.) wandering rapidly over ground in open, grassy-weedy areas; on
numerous low-growing, herbaceous plants, including *Arctotheca calendula and
*Echium lycopsis, etc. (dets. NM). These larvae are fairly common here, but far
less abundant than those of S. glatignyi, which occur in exactly the same places at
the same season. The Amsacta larvae are quickly identified by their more rapid
locomotion and less-dense setae; univoltine here, with adults emerging in autumn
(+). See Common (1966b: 72, 111) for color illus. and photo (B. & W.) of adult.
(Preserved = 1, 5, 6.)
Ardices—see Spilosoma.

e Spilosoma canescens Butl. (det. NM)—Ar.41. S.AUST., higher parts of the
Mt. Lofty Range, Aldgate district (NM): Captive larvae (Jan.) readily accepted
numerous offered plants (both herbaceous and woody). Three woody plants for
which they showed a great liking were lvs. of ash, *Fraxinus sp—OLEACEAE;
apple, *Malus sp.—ROSACEAE;; pine, *Pinus sp.—PINACEAE (dets. NM). Pinus,
in particular, was eaten avidly! Adults (local; B—) fly Nov.—Jan.; in 5 yrs. never
recorded at Blackwood or other lower elevations near Adelaide. Probably univoltine.
See Common (1970: 857) for adult photo. (Preserved = 1-4, 9.)

e Spilosoma glatignyi (LeGuill.) (det. IC)—Ar.34 & 34A. (1) S.AUST., Black-
wood-Belair-Eden Hills (NM): Larvae (June—mid Oct.) exceedingly common most
years, primarily on various low-growing herbaceous plants or shrubs, often becoming
pests in gardens. Some of the “primary” foodplants: *Echium lycopsis, *Genista
maderensis, Pultenaea largiflorens var. latifolia, *Arctotheca calendula, *Chrysanthe-
moides monilifera, Olearia ramulosa, and *Plantago lanceolata (dets. MK). Of the
above, the first two listed are perhaps the most important foodplants of this moth
around Blackwood. In addition to the above-listed, I have seen them feeding on
numerous other plants, both native and naturalized; univoltine. Adults (A+) fly
mid Jan._mid May (peak April); the majority come to uv. light mostly after 2300
hrs. See Common (1966b: 111) or Tillyard (1926: Pl. 33) for adult photo. (Pre-

18 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

served = 1, 4-6, 8dd, 9; photos = Ic, 6.) (2) S.AUST., S coast of Kangaroo Is.,
at Seal Bay (NM, M. Pate, & G. D. Seton): Fullgrown larvae (16 Oct. 66) crawling
over the sand, in association with larvae of my An.6A (Anthelidae) and Lp.12
(Lymantriidae); seen eating the same herbaceous plants as listed for the latter.

e Utetheisa lotrix (Cram.) (det. NM)—Ar.36. N.TERR., 15 mi. S of Mataranka,
at Warlock Ponds (NM): Larvae (9 April 66) on lvs. and floral parts of Crotalaria
trifoliastrum Willd—FABACEAE (det. NM). See Common (1966b: 111) for
adult photo. (Preserved = 4-6, 9.)

e Utetheisa pulchelloides Hamps. (det. SF)—Ar.35, 35A, & 35B. (1) S.AUST.,
Blackwood and Eden Hills, on open & dry-grassy slopes (NM & J. Herridge): Larvae
(9 Jan. 66) commonly feeding on floral parts of *Echiuwm lycopsis (det. NM).
Adults (A+) fly Sept.-mid May (peaks Dec.—Feb. & April). (Preserved = 1, 4-6,
8d, 9.) (2) S.AUST., 4 mi. E of Two Wells, in dry, weedy-grassy paddocks and at
roadsides (NM, TN, & G. Furness): Larvae (19 March 67) exceedingly common on
potato weed, *Heliotropium europaeum 1.—BORAGINACEAE (det. MK). (3) S.
AUST., Adelaide suburb of Heathpool (R. Edwards): Larvae (30 April 67) in a
garden, defoliating the ornamental forget-me-not, *Myosotis sp—BORAGINACEAE
(det. NM).

(B) SuBFAMILY LITHOSIINAE

e Palaeosia bicosta (Walk.) (det. SF )—Ar.43. S.AUST., Mt. Lofty Range, Belair
Nat. Park, nr. Waverley Lodge (NM & D. Bakker): Larvae (13 Sept. 69) on stems
or branches of various woody native shrubs, especially Acacia pycnantha, feeding on
LICHENS (and other minute plant growths, probably algae) that grow on the often
rain-wet stem surfaces in winter and early spring. See Common (1966b: 111) for
adult photo. (Preserved = 1, 4-7.)

e Scoliacma bicolora (Bdv.) (det. NM )—Ar.37. S.AUST., Blackwood, Hannaford
Rd. (NM): Larvae (July—early Oct.) on the open or bare ground of trails through
thickets of Acacia pycnantha and Eucalyptus odorata, or on + bare patches of firm
ground, where they wander slowly over the winter-damp surface “grazing” on an
abundant MOSS(!), Pottia sp—POTTIACEAE (det. A. Mitchell). They are diurnal
and especially crepuscular feeders, active on mild or warm days, either cloudy or
sunny (if the sun is not intense or constant). Occasional individuals have also been
seen eating one of the lichens, especially where these are growing on large rocks, in
open but densely grass-covered paddocks (Belair district). But this common moss
appears to be by far the primary foodplant around Blackwood. One larva (in
captivity) was also seen eating an unidentified liverwort. Adults (B+) fly Oct.—
Nov. (peak); again (B—) from Jan.—mid April. Individuals of the summer—autumn
brood are generally of smaller size and less intense coloration. See Common (1966b:
72) or Tillyard (1926: Pl. 27) for adult in color; McFarland (1970: 350 & 1972b:
229) for egg photos. (Preserved = 1-7, 8dd, 9; photos = 1, 2, 5-7.)

Thallarcha—see Xanthodule.

e Xanthodule ombrophanes (Meyr.) (det. IC)—Ar.38. S.AUST., Blackwood-
Belair districts (NM): Larvae (Sept.-March) on stems or branches of numerous
woody plants, where they feed along the stem surfaces, probably on lichens and/or
algae (exact food sources not positively identified). They are most often collected
by searching over the branches of Bursaria spinosa and Casuarina stricta, or by
beating C. muelleriana and Exocarpos cupressiformis; also generally encountered
when beating most of the other medium-sized or larger native shrubs of this habitat.
In no instance has a captive larva of this moth ever been observed feeding upon the
leaves (or floral parts) of any of the numerous plants upon which they have been
collected. The newly-hatched larvae have first instar dispersal behavior (see Mc-
Farland, 1973: 205-207). Adults (6 ¢ only; B+) fly Oct._June (peaks Oct.—Nov.;
late Jan.—early Feb.; April-May), coming to uv. light esp. after 2300 hrs. Sexual
dimorphism is stupendous! The flightless (brachypterous) @ waits on its cocoon

SUPPLEMENT TO VOLUME 33 19

for arrival of the ¢ (as in N. American lymantriids: Hemerocampa); after mating,
all its eggs are deposited, without any type of covering, on the cocoon surface. For
adult 2, cocoon, and egg photos of this sp., see McFarland (1970: 350 & 1972b:
229); also Common (1966b: 109) for larval photo of a related sp. (Preserved =
eee —7, 9: photos= ¢ 21, Plc 2 2c, 2Ec, 7, Tc.)

(C) Susramity NycTEMERINAE

e Nyctemera amica (White) (det. IC)—Ar.39 & 39A. (1) S.AUST., Blackwood
(NM): Larvae (especially summer) on the S. African herbaceous, perennial vine,
Cape “ivy, *Senecio mikanioides Otto ex Walp —ASTERACEAE (det. NM). Adults
(B) recorded for all months, with peaks in Jan—March & May-June; both diurnal
and nocturnal activity, coming to uv. light esp. after 2300 hrs. See Common (1966b:
113 or 1970: 857) for ¢ adult photo; McFarland (1970: 350 & 1972b: 229) for
egg photos. (Preserved = 1-6, 8dh, 9; photos = 1 pair in cop., 2, 4-6.) (2) S.
AUST., nr. tip of Yorke Peninsula, + 2 mi. NW of Jim Brown’s Well (NM, N. B.
Tindale, & P. Aitken): Larvae (3 Nov. 65) on the native Senecio aff. lautus Forst.
f. ex Willd. (det. MK). (Preserved = 5.)

e Nyctemera baulus (Bdv.)—see Supplement on Fiji (following Zygaenidae).

e Nyctemera secundiana Lucas (det. IC)—Ar.40A. N.QLD., Atherton Tableland,
12 mi. NE of Atherton, at Tinaroo Pines Caravan Park (+ 2500’) (D. & NM):
Larvae (April-May 72) on the soft, rank-growing Asian weed, known as thick-head,
*Crassocephalum crepidioides (Benth.) J. Moore—ASTERACEAE (det. NM).
Adults both diurnal and nocturnal. (Preserved = 1-6, 9.)

CARTHAEIDAE

© Carthaea saturnioides Walker (det. NM)—Ca.1, 1A, 1B, 1C, & ID. (1) W.
AUST., Stirling Range, along roadside, nr. Toolbrunup Peak, + 200 mi. SE of Perth
(NM &N. B. Tindale): Larvae abundant (19 Nov. 68), along with a few unhatched
eggs, on young lvs. of the low sclerophyll shrub, Banksia sphaerocarpa R. Br. (det.
WAH). None of the vivid red mites, found so abundantly on my An.10 ( Anthelidae),
were seen on these larvae, although they were often closely associated, on the same
bushes, with the (less numerous) An.10 larvae. Adults in this locality probably
fly from Oct.—early Dec.; one fairly worn ¢ was taken at uv. light betw. 0300-0400
hrs on 20 Nov. 68; although the light was running all night, none arrived before 0300
hrs; univoltine. Duplicate copies (8 pp.) of detailed field and larval behavioral notes
on this superb moth were deposited in the Lepidoptera Section of the A.N.I.C.
(1968). See Common (1966a; 1970: 850, 858) for illus. of adult, larva, & pupa;
McFarland (1970: 349 & 1972b: 225) for egg photos. (Preserved = 2-4, 9; photos =
1, Ic, 2, 4, 4c, 5, 5c, 6.) (2) W.AUST., 1 mi. W of Needilup (NM & N. B. Tindale):
Larvae, mostly last instar & penult. (22 Nov. 68), abundant on young lvs. of the
tough-sclerophyll shrubs, Dryandra cirsioides Meisn. and (to a much lesser extent)
Banksia caleyi R. Br—both PROTEACEAE (dets. WAH); the former appears to be
the primary foodplant in this locality. (Preserved = 1, 5, 6, 9; photos = 1, lc, 5, 5c,
6.) (3) W.AUST., 10 mi. E of Jerramungup (NM & N. B. Tindale): One Ist
instar larva (24 Nov. 68) found feeding on the brown-pubescent young lvs. of
Dryandra pteridifolia R. Br. (det. WAH). (There was no D. cirsioides anywhere
nearby.) (Preserved — 3.) (4) W.AUST., 49 mi. E of Ravensthorpe, 1 mi. W of
Munglinup (NM & N. B. Tindale): Last instar larvae (24 Nov. 68) feeding on
young lvs. of D. pteridifolia and on one other unident. low-growing Dryandra sp.,
close to (but smaller than) pteridifolia. (D. cirsioides was common here but no
larvae, or their unmistakable pinkish frass, could be found on it during a 30 min
search by both of us.) (5) W.AUST., 9 mi. S of Ravensthorpe, on road to Hope-
toun (NM & N. B. Tindale); Two 3rd instar larvae found (25 Nov. 68), on two
widely-separated individuals of the linear-leafed sclerophyll shrub, Grevillea concinna

20 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

R. Br. (det. WAH), eating young lvs. only. This plant was in bloom at the time
(deep red fls.). Dryandra cirsioides was more abundant than G. concinna at this
location, but no larvae, frass, or evidence of feeding could be located on the D.
cirsioides bushes. (6) W.AUST., Moresby Ranges, Ooakajee-Howatharra-Nanson dis-
trict, between 15-25 mi. NE of Geraldton; esp. in and nr. Howatharra Hill Reserve
(McFarland, 1977: 19): Larvae fairly common some years (Sept._mid Oct.) on
tender new lvs. (only) of Dryandra fraseri R. Br. (det. WAH); this is a dense, low-
growing, extremely sclerophyll sp. with very tough and prickly mature lvs.; it grows
mostly on brown lateritic gravel. Adults probably fly from about mid Aug.—early
Oct. here.

COCHYLIDAE (PHALONIIDAE)

e Hyperxena sp. (det. IC)—46(M). W.AUST., Moresby Range, Red Peak district,
+ 13 mi. NE of Geraldton (D. & NM): Larvae (2 Aug. 73) extremely abundant on
the small, pink fls. & fl. buds of the large shrub, Scholtzia Pparviflora F. Muell.—
MYRTACEAE (det. NM). The habits and behavior of these larvae are strongly
reminiscent of lycaenids, as is their general appearance at first glance. The macula-
tion is a colorful blending of pink and white, which is superbly cryptic on the
foodplant inflorescences. They usually rest tightly curled around the fls. and bud
clusters, almost invisible if not moving, but are easily discovered by beating if present.
Adults emerged mid June 74. (Preserved = 1, 4—7, 8h, 9. )

COSSIDAE

e Ptilomacra senex Walk. (det. NM)—Co.29(M). S.AUST., Mt. Lofty Range,
in Belair Nat. Park nr. Sheoak Rd., 1 mi. E of Belair railway station; also around
Montacute, and in other localities from NE to S of Adelaide (NM): Egg masses
(mid Aug.—Nov.) often found attached to the stems of various low-growing plants
or bushes, including the long lvs. of Xanthorrhoea semiplana FvM. (det. NM), which
I strongly suspect is the Belair locality foodplant. (The larvae are presumably
borers.) This highly-conspicuous egg mass could not be mistaken for any other;
it is composed of large numbers of very large, grayish-black eggs, tightly glued to
each other with a shiny adhesive, and usually deposited in long encircling-bands.
Adult sexual dimorphism and first instar larval dispersal are notable in this species.
See Common (1966b: 27) for ¢ adult photo. (Preserved = 2, 3.)

GEOMETRIDAE

Among the geometrids listed here (particularly ennomines), reference is frequently
made to a “rain-hatching tendency” in the eggs of certain species, or to “first instar
dispersal.” These phenomena were described and discussed at some length
earlier (McFarland, 1973: 203-206), so will not be repeated here. That paper was
intended to directly follow a series of egg photographs published earlier (1972b),
to which it frequently refers, but publication was unexpectedly delayed and several
items were accidentally omitted after correction of the proofs; these are included as
annotations in the list that follows.

(A) SuBFAMILY ENNOMINAE (SYNONYM = BOARMIINAE )

Amelora—see also Diastictis and Loweria.
e Amelora crypsigramma Lower (det. IC, NM). W.VIC., 5 mi. S of Kiata, Lowan
Reserve (NM, collector): Larva (28 Sept. 67) on the woody shrub, Baeckea behvrii
(Schldl.) FyVM.—MYRTACEAE (det. MK). Adult @ emerged 3 April 68. (Pre-

served = 1, 6.)

SUPPLEMENT TO VOLUME 33 91

e Amelora fucosa Turner (det. IC)—G.183 & 183A. (1) S.AUST., nr. Port Wake-
field, on top of Hummock Mt. in dense scrub (H. M. Cooper): Larvae (31 Aug. 67)
on Beyeria sp—EUPHORBIACEAE (det. NM). (Preserved = 5, 9; photos = 5.)
(2) S.AUST., just W of Alawoona, at roadside (D. & NM): Larvae (late July 71)
on the low, woody shrub, Beyeria opaca FvM.—(det. NM). 4 adults (all ¢ 2)
emerged late March-mid April 72; probably univoltine. Considerable variation is
shown in forewing groundcolor, but not in maculation. (Preserved = 1, 4, 5, 6, 9;
photos = 5.)

e Amelora leucaniata Gn. (det. SF)—G.171. S.AUST., Mt. Lofty Range, Black-
wood (NM): Captive larvae (June—Aug.) readily accepted lush young lvs., buds,
& fls. of the annual weed, *Polygonum aviculare L.—POLYGONACEAE (det. NM);
a preference was shown for the large-leafed, luxuriant young seedlings of this plant,
which are well started (2” high +) by early July. Adults (C) fly March—April;
those reared in captivity emerged in Dec. (possibly not true to natural conditions
here or possibly representing an early summer brood?) The eggs are rain-hatched.
(Preserved = 1-6, 9; photos = 1, 2, 5, 5c, 6.)

e Amelora macarta Turner (det. IC)—G.151, 151A, 151B, & 151C (under A.
petrochroa Guest and A. poliophara Tumer, in some collections). (1) S.AUST.,
+ 8 mi. NE of Two Wells, at edge of sandhills (NM & TN): Larvae (20, 27 Aug. 66)
only on the twining parasite, Cassytha pubescens, which was growing in shrubs of
Calytrix Pinvolucrata J. M. Black (dets. MK). Four adults emerged April 67. (Pre-
served = 1, 4-6, 9; photos = 1, 5c.) (2) S.AUST., Mt. Lofty Range, Belair National
Park, 34-1 mi. E of Belair railway station (NM); also, 4 mi. S of Ashbourne (J. O.
Wilson & NM): Larvae (July—Aug. 68) only on Cassytha pubescens, which was (in
these localities) mostly parasitizing Leptospermum myrsinoides, Casuarina muel-
leriana, and Olearia ramulosa (dets. NM); also, in one instance, parasitizing
Exocarpos cupressiformis and harbouring many larvae of A. macarta. The last instar
larvae show various maculation-phases, one being very strongly marked with 9 or 10
dark, oblique dorso-lateral lines, and the others partially or even totally lacking these
prominent marks; all forms were regularly encountered in Belair Nat. Park. Adults
emerged March—April 69; univoltine. (Preserved = 1, 5, 6, 9; photos = 1, 5,6.) (3)
W.AUST., + 16 mi. N of Geraldton, nr. Oakajee (D. & NM): 2 last instar larvae
(28 July 72) on Cassytha sp., nr. glabella R. Br., which was parasitizing Casuarina
campestris Diels. (dets. NM). Adults emerged mid-late April 73. This may
represent a distinct subspecies. Also common at Drummond Cove, W. A.; larvae on
Cassytha sp., nr. pubescens R. Br., during June and July. (G.151C: Preserved =
1-6. 9.)

e Amelora milvaria Gn. (det. IC, NM)—G.114. S.AUST., Blackwood-Belair district
(NM): Larvae (July—Sept.) primarily on lvs. of Pultenaea largiflorens var. latifolia,
young lvs. of Acacia pycnantha, and mature (tough) lvs. of Leptospermum myr-
sinoides; also (less often) seen feeding on young lvs. of Acacia myrtifolia (Sm.)
Willd., fls. of Hakea rugosa R. Br., foliage of Exocarpos cupressiformis, lvs. of
Calytrix tetragona, young lvs. of *Eucalyptus ficifolia F. Muell. (in a garden), lvs. of
Olearia ramulosa, and new lvs.-buds-fls. of *Chrysanthemoides monilifera (dets. NM).
Adults (A) fly mid March-early May. The eggs are rain-hatched. (Preserved =
1-6, 8h, 9; photos = 1, 5c.)

e ?Amelora sp. (det. NM)—S.AUST., Mt. Lofty Range, + 4 mi. S of Ashbourne
(J. O. Wilson): 2 larvae (23 Aug. 68) on the dense-shrubby Casuarina muelleriana
(det. NM). Adults emerged 4 April (4), and 6 April 69 (2). This uncertain
determination is included because of the very distinctive adults and the interesting
foodplant record; it is hoped that more specimens may eventually be taken. The 2
reared adults are in the A.N.I.C., where the late J. O. Wilson placed his collection
- before he died in 1972. (Preserved = 1, 6; photos = 1.)

bo
bo

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Angelia—see Loweria.
e Boarmia cognata Walk. (det. IC, SF)—G.91. S.AUST., Blackwood-Belair district
(NM): Larvae (most months) common on foliage of the shrubby root-parasite,
Exocarpos cupressiformis (det. NM). Adults (B+) recorded for every month (peaks
in Dec.—Jan. & April). (Preserved = 1, 4—6, 8h, 9; photos = 1, le, 2, 5.)
e Boarmia Ploxographa Turner (det. SF, NM)—G.115. (1) S.AUST., Blackwood-
Belair district (NM): Larvae (July—Oct.) on many native shrubs, including (par-
ticularly) Pultenaea largiflorens var. latifolia and Bursaria spinosa (dets. MK); also
seen feeding on young lvs. of Acacia pycnantha, Exocarpos cupressiformis, Olearia
ramulosa, and *Chrysanthemoides monilifera (dets. MK). Adults (A) fly spring—
autumn (peak March—April); sexual dimorphism is fairly pronounced. (Preserved =
1, 5, 6, 9: photos = 1, 2.) (2) S.AUST., Hallett Cove, on@ecastalilutiisn Sor
Adelaide (NM): Larvae (July—Oct.) fairly common on the dominant low-woody
shrub, Beyeria leschenaultii var. latifolia Grining—EUPHORBIACEAE and _ the
woody shrub, Alyxia buxifolia R. Br.—APOCYNACEAE (dets. MK). No adults
were reared, so identification is not absolutely certain; these larvae appeared identical
to those of the Blackwood population, however. (Preserved = 5, 8hh, 9; photos = 5.)
e Boarmia penthearia Gn. (det. NM)—G.211 (syn. = rupicolor Butl.). (1) S.
AUST., Blackwood (NM): Captive larvae (March 70) readily accepted young
phyllodes of Acacia pycnantha (det. NM). Adults (C) fly summer—autumn. First
instar larval dispersal is notable in this species. (Preserved = 1-6, 9; photos = 1, 2.)
(2) W.AUST., Drummond Cove, + 7 mi. N of Geraldton (NM): Larvae (spring—
summer) on Acacia ligulata A. Cunn. ex Benth. (det. B. Maslin). Adults (B) fly
most months.
@ Boarmia suasaria Gn. (det. IC, SF)—G.78. S.AUST., Blackwood-Belair district
(NM): Larvae (most months, with a peak in summer) on foliage of only the tree,
Casuarina stricta (det. NM); never found on other (shrubby) Casuarina spp. fre-
quently beaten in this locality. Adults (B+) fly Dec.—mid July (peaks Jan. & late
March—April). First instar larval dispersal is notable in this species. (Preserved =
1-6, 9; photos = 1, 2, 5.)
@ Boarmia sp. (det. NM)—G.215. S.AUST., 4 mi. E of Two Wells, Lewiston Park
(NM & D. Bakker): Larvae (4 Sept. 69) on the tree, Callitris preissii Miq.—
CUPRESSACEAE (det. MK). This incomplete determination is included because
of the interesting foodplant record; the adult is distinct from all foregoing spp.
(Rreserviedi— lp son)
e “Capusa” chionopleura Turner (det. IC, NM )—G.230. (This unique moth is in
no sense referrable to Capusa, but apparently no other genus is available for it at
present; nor does it show close kinship with either G.231 or 237.) | W.AUST., Drum-
mond Cove, +7 mi. N of Geraldton (D. & NM): Captive larvae (July—Aug. 72)
readily accepted only the (orange) fls. and fl. buds of the leafless intricate shrub,
Daviesia divaricata Benth— FABACEAE (det. NM). Adults (D) fly June-July;
more common inland, or further north in more arid localities; univoltine. (Preserved
== 1-6) 9 photos leon ocx)
© Capusa senilis Walk. (det. IC)—G.235. W.AUST., Drummond Cove, +7 mi.
N of Geraldton (D. & NM): Captive larvae (July—Aug.) readily accepted phyllodes
of Acacia ligulata A. Cunn. ex Benth. (det. B. Maslin); also one record (10 Sept.
76) of a healthy last instar feeding on lIvs. of the edible garden pea, *Piswm—
FABACEAE. The latter had probably been on the pea vine all its life, as much
evidence of past and recent feeding surrounded the area where it was at rest, and
no other possible foodplant was growing anywhere nearby. It was collected and it
continued to feed well (for several more days) on the pea leaves, until the pre-pupal
wandering stage arrived. A. ligulata is undoubtedly the usual foodplant in this
locality; new growth is preferred if available. These larvae (and the eggs) show
much closer affinities with G.105 than with G.231 or 237; clearly, neither of the
latter belong in the same genus with senilis and stenophara. Adults (B+) fly May—

SUPPLEMENT TO VOLUME 33 23

July (peak late June-early July); univoltine. Regarding adult Capusa spp. resting
position, see p. 13. (Preserved = 1-6, 9; photos = 1, 5, 5c.)

@ Capusa stenophara Turner (det. IC)—G.105. (This is a synonym of cuculloides
Felder according to evidence from genitalic slides prepared and studied by SF,
April 71.) (1) S.AUST., Blackwood (NM): Captive larvae (Aug.—Sept.) readily
accepted young lvs. of Eucalyptus odorata; larvae (in the field) often found on
young lvs., fl. buds, and fls. of Acacia pycnantha, as well as Eucalyptus odorata
(dets. NM). Adults (B) fly late May—July (peak mid June); activity primarily
from dusk to + 2100 hrs (rarely later), often on cold (38—45°F) and clear nights
when almost nothing else is on the wing; univoltine. First instar lerval dispersal is
notable in this species. See McFarland (1972b: 239) for egg photos. (Preserved =
1-6, 9; photos = 1, 2, 5, 5c, 6.) (2) S.AUST., Happy Valley and Aldinga scrub,
S of Adelaide (Mr. and Mrs. J. O. Wilson): Larvae (Aug.—Sept. 67) on fl. buds
& fls. of Acacia pycnantha (det. NM). (3) W.VIC., 5 mi. S of Kiata, Lowan
Reserve (NM): Young larvae (28 Sept. 67), of an unidentified Capusa sp., beaten
from the upright woody shrub, Melaleuca uncinata R. Br. ex Ait., in close assoc. with
Baeckea behrii (Schldl.) FvM.—both MYRTACEAE (dets. MK); the circumstances
indicated the former as the probable plant from which these larvae fell.

®@ “Capusa” sp. (det. NM )—G.231 & 231A. (This moth is probably undescribed; it is
fairly near Capusa, but clearly belongs in another genus; the larvae show closer
affinities with G.237 than with typical Capusa larvae. ) GRY WEAUSE. “SE o£
Onslow, at Nanutarra (D. & NM): Original 2 moth collected at light; eggs obtained.
Captive larvae (July—Aug. 72) reared at Drummond Cove, W.AUST., where they
readily accepted young phyllodes of Acacia ligulata A. Cunn. ex Benth. (det. B.
Maslin). (Preserved = 1-6, 9.) (2) W.AUST., Drummond Cove, +7 mi. N of
Geraldton (D. & NM): Captive larvae (June-July ) readily accepted young phyllodes
of Acacia ligulata A. Cunn. ex Benth. (det. B. Maslin). Adults (A) fly late May—
early Aug. (peak late June—early July); univoltine. (Preserved = 1-6, 9; photos =
ie oc:)

® “Capusa” sp. (det. NM )—G.237. (This distinctive moth is probably undescribed; it
is fairly near Capusa, but clearly belongs in another genus; the larvae show closer
affinities with G.231 than with typical Capusa larvae.) W.AUST., Drummond Cove,
+ 7 mi. N of Geraldton (D. & NM): Captive larvae (mid July—mid Sept.) readily
accepted fl. buds & fls. (only) of Acacia ligulata A. Cunn. ex Benth. (det. B. Maslin);
these larvae are clearly adapted, both in behavior and coloration, specifically to
blossom-feeding on acacias. Adults (B—) fly June-July (peak late June). (Pre-
served = 1-6, 9; photos = 1, 5, 5c.)

e@ Casbia lithodora Meyr. (det. SF, NM)—G.160. (1) S.AUST., Yorke Peninsula,
+ 2 mi. S of Kainton (NM, N. B. Tindale, & P. Aitken): Single larva (4 Nov. 65)
on the open-airy shrub, Pomaderris paniculosa FvyM. ex Reisseck—RHAMNACEAE
(det. MK), along with many larvae of G.131; adult 2 emerged Dec. 65. (Preserved
= 1, 6.) (2) S.AUST., at base of Black Hill, Athelstone (Addison Ave.), E of
Adelaide (NM & TN): Larvae (22 Oct. 66) common on young and mature lvs. of
the woody dwarf shrub, Cryptandra tomentosa Lindl—RHAMNACEAE (det. MK).
Adults fly late spring-summer; they are easily alarmed to flight in the daytime
(observation of TN, 15 Jan. 67), but are probably not truly diurnal. (Preserved =
1, 4-7, 8h, 9; photos = 1, 5c.) (3) W.AUST., 5 mi. N of Geraldton (Glenfield
district), Beatie Rd., Lot 131, in an Acacia-Banksia association (D. & NM): One
larva (mid Nov. 72) on Cryptandra sp. (det. NM); died in last instar (not pre-
served), but identical to well-marked larvae of the Black Hill population. (SF,
in a letter of 8 July 1971, stated that there are “several from Geraldton and one from
Perth” in the B.M.(N.H.) series of adult specimens. )

®@ Casbia ?rhodina Turner (det. IC)—G.131. S.AUST., Yorke Pen., + 2 mi. S of
Kainton (NM, N. B. Tindale, & P. Aitken): Larvae (4 Nov. 65) fairly common on
Pomaderris paniculosa FvM. ex Reisseck—RHAMNACEAE, (det. MK); they match

94 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

to perfection (in color and maculation) the undersides of the foodplant lvs. Adults
emerged late spring-summer. (Preserved = 1, 5-7, 8h, 9.)

Ceratucha—see Ciampa.

“Chlenias’—see also Ciampa and “Miscellaneous Unidentified Ennominae” at
end of this subfamily.
e “Chlenias” melanoxysta Meyr. (det. IC & NM)—G.221 (The type-specimen is
Chlenias melanoxysta Meyr., in the S. Aust. Museum; an apparent synonym is the
oenochromine, Cycloprorodes apalama Turner. Clearly, melanoxysta does not belong
in Chlenias.) S.AUST., Belair, at 129 Gloucester Ave.; rarely taken at Blackwood;
probably more common between Kalyra Rd. and Windy Point, where Dodonaea is
abundant (?) (D. & NM): Captive larvae (June-July 71) readily accepted lvs. of
(only ) Dodonaea viscosa (det. NM), esp. the new growth. These larvae are difficult
to rear, needing abundant daily moisture combined with excellent ventilation and
low temperatures. They show not the remotest kinship with Chlenias larvae. Adults
(C) fly only mid May-mid June (peak late May); univoltine. The eggs seem to
be on the borderline of a rain-hatching tendency. (Preserved = 1-4, 6, 9; photos =
Jie Mien, 2ut5e)
e Chlenias ?pachymela Lower (det. NM)—G.106 & 106A. S.AUST., Blackwood-
Belair district (NM): Larvae (July—Oct.) on a wide range of native and introduced
woody plants, some specific examples being: Bursaria spinosa, Dodonaea viscosa,
young lvs. of Acacia pycnantha, Pultenaea largiflorens var. latifolia; one record on
a plum tree, *Prunus sp., and one on young lvs. of loquat, *Eriobotrya sp.—both
ROASACEAE; and one feeding well on fl. buds of *Camellia sp.—THEACEAE
(dets. NM). Of the native flora in this locality, the first 3 plants listed appear to
be preferred, particularly Bursaria. These larvae always have a distinctive raspberry-
red blotch (which fades out completely in alcohol) near the base of the A6 proleg;
the head is notably bulbous, light golden-brown, and only faintly-marked. Disturbed
fullgrown larvae are usually reluctant to release hold or drop; if dropping, usually
on a tough silk thread. Adults (B+) fly mid May—July, and are the largest (and
“heaviest’) of the Chlenias spp. in this locality; univoltine. The eggs are rain-
hatched. (Preserved == 1-7, 9; photos = 1.)
e “Chlenias” rhynchophora Lower (det. SF, NM)—G.127 & 127A. (The @ type
specimen is Chlenias rhyncophora Lower, in the S.Aust. Museum. This moth
clearly has no kinship with Chlenias; it appears to belong in the genus Rhynchopsota. ).
S.AUST., Belair Nat. Park, 1 mi. E of Belair railway station; also the Eden Hills-
Blackwood district (NM & TN; G. Furness): In the first locality larvae (Oct—early
March) are mostly on the locally-abundant shrub, Casuarina muelleriana, but also
on the tree, C. stricta (dets. MK). In the two latter localities, they are mostly on
C. stricta, as C. muelleriana is extremely rare here. There is a major population of
this sp. below 5 Yalanda St., Eden Hills. In common with a number of Australian
oenochromine genera (many Dichromodes spp., Monoctenia, and Phallaria are ex-
amples), the larval growth-rate in rhyncophora is very slow under natural conditions;
interestingly, they are on the plants throughout the hottest and driest months of
the year (Nov.—March), a time when most univoltine spp. are in pupal diapause
here. The eggs take 6-7 weeks to hatch, and the larval stage lasts a full 8 months
(about mid July—mid March)! This is altogether an aberrant species in several re-
spects. The larvae (also the eggs and pupae) show no affinities with Chlenias spp.,
but do appear to be fairly close to the Dichromodes larval type with regard to gen-
eral appearance, profile, growth-rate, habits, and behavior. Adults (A, nr. 5 Yalanda
St., Eden Hills; B+, in most parts of Blackwood) fly mid May—mid July (peak
only late May—mid June); univoltine. See McFarland (1972b: 237) for egg photo.
(Preserved = 1-6, 9; photos = 1, 2, 4-6.)
e Chlenias seminigra Rosenstock (det. NM)—(G.220 ?). S.AUST., Mt. Lofty
Range, Aldgate (Heather Rd.) (NM & D. Bakker): (a) Mature larva coll. by
K. Sandery (Sept. 70) on the parasitic twiner, Cassytha sp. (det. K. Sandery);
6 adult emerged 1 June 71. (b) Mature larva (13 Sept. 69) on fl. buds and fls.

SUPPLEMENT TO VOLUME 33 25

of the dwarf shrub, Dillwynia hispida (det. NM). I suspect that the single larva
collected by Sandery (a), which I never saw, and my single larva (b), for which
no adult was reared, are one and the same species. The fullgrown larva (b), my
G.220, is deposited in the S. Aust. Museum, and the reared adult (a) is in the
K. J. Sandery collection. The G.220 larva shows strong affinities with my G.111, but
is somewhat larger and is more extensively and colorfully-marked. These incomplete
records are included because of the apparent rarity (or very localized occurrence)
of C. seminigra in this part of South Australia. The dark brown to sooty-tinged adult
is quite unlike any other Chlenias here discussed. Probably univoltine. (Preserved
—wole(o:). 6; photos = ¢1.)

e Chlenias sp., near stenosticha Turner (det. IC, NM)—G.111, 111A, & 239.
(1) S.AUST., Blackwood-Belair district (NM): Larvae conspicuous (July—Sept. )
on fl. buds, fls., and young growth of various native woody shrubs, esp. Cryptandra
tomentosa Lindl.—RHAMNACEAE and Grevillea lavandulacea; sometimes on buds
of Dillwynia hispida and young lvs. of Calytrix tetragona (dets. MK). These larvae
appear to be primarily bud and flower feeders in this locality, if allowed their
preference. Very colorful: among the distinctive markings are two intense reddish-
brown lateral spots, one each on T2 and T3, just above the broad and prominent
yellow-cream spiracular line, which extends the full length of the body. Fullgrown
larvae are quick to curl up and fall free (never on a silk thread) if disturbed. Adults
(B—) fly April-May (univoltine); they are rather frail when compared to other
Chlenias spp. occurring here, and are the smallest sp. around Blackwood; although
uncommon at my lights, the larvae of this sp. were more frequently encountered
here (1965-1970) than were those of G.112. (Preserved = 1-6, 9; photos = I, 5.)
(2) S.AUST., nr. Hallett Cove railway station (NM): 2 small larvae (6 Aug. 67)
on fl. buds and fls. of Acacia rotundifolia Hook. (det. NM). (3) W.AUST.,
Drummond Cove, +7 mi. N of Geraldton (D. & NM): Captive larvae (June-—early
Aug. 73) readily accepted new and semi-mature phyllodes of Acacia ligulata A.
Cunn. ex Benth. (det. B. Maslin). Growth was rapid and disease-free on this plant.
Although initially code-numbered G.239, it became evident by last instar that these
larvae merely represented a western population of my South Australian G.111 ma-
terial. (Preserved = 1-6, 9.) (4) W.AUST., in Spalding Park, Bluff Point district,
N of Geraldton (D. & NM): Larvae (July, 1973) on fls. and new lvs. of Grevillea
pinaster Meisn. (det. WAH). (Preserved = 5.)

@ Chlenias ?umbraticaria Gn. (det. SF, NM)—G.112. (1) S.AUST., Blackwood-
Belair district (NM): Larvae conspicuous (July—Sept.) on fl. buds, fls., and young
growth of various native woody shrubs, quite often in association with larvae of the
foregoing sp. (G.111). Favoured foodplants here are fls. of Hakea rostrata and fls.
of Cryptandra tomentosa Lind|—RHAMNACEAE; also (less often) on fils. of
Grevillea lavandulacea, fls. of Pultenaea largiflorens var. latifolia, or on Cassytha
pubescens (dets. MK). Fullgrown larvae of this moth are similar, in both habits
and behavior, to those of G.111, but are entirely different in details of color-
maculation; they are also “heavier” (more massive) in the whole body. Very
colorful: some of the distinctive marking are about 10 well-separated brick-red (or
orange-brown ) elongate lateral marks, one per segment, on the white spiracular line
(from T2—A8); dorsum and sides (above spiracular line) entirely without other
major stripes, the whole area being reticulate-speckled in a pattern of dark gray-
brown rings, surrounding whitish dots, which creates an overall effect of being
heavily and uniformly “salted” on a dark background. Adults (B) fly late March—
April (univoltine); they are quite unlike any of the other Chlenias spp. here listed,
and (atypically for this genus) show only minor individual variation in color and
maculation. The eggs are rain-hatched. (Preserved = 1-6, 8h, 9; photos = 1, 2, 5,
de, 6.) (2) W.AUST., +5 mi. N of Geraldton (Glenfield dist.), along Beatie Rd.
(D. & NM): One larva (20 July 73), of what is almost certainly this sp., found
among low shrubs at roadside, resting in full view, head downward; foodplant
unidentified. (Preserved = 5.)

26 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

e Chlenias sp. nov., near banksiaria LeGuill. (det. SF)—G.103, 103A & 103B. S.
AUST., Blackwood-Belair district and Hallett Cove (NM): Larvae (June—Sept.)
on several native woody shrubs; by far the preferred foodplant (of the Blackwood
population) appears to be Olearia ramulosa, which bushes they nearly defoliated in
some years during the winter (along Hannaford Rd.); they were often abundant on
both Dodonaea viscosa and O. ramulosa at Windy Point (Belair); along Gloucester
Ave. (Belair), and in parts of Blackwood, they were sometimes common on Bursaria
spinosa; occasionally also on *Chrysanthemoides monilifera and *Genista maderensis
(dets. NM). At Hallett Cove, I found them only on Olearia ramulosa (fairly com-
mon, but never in the great numbers sometimes seen at Blackwood). Although
colorful, they blend in fairly well with the “intricate confusion” of shadows and
thin (linear) O. ramulosa lvs.; the overall pattern on the larva (dorsolaterally) is
of many fine parallel lines of black, white, and yellow to orange. The single strong
middorsal line is pale to deep orange in living larvae of this species, and on that
alone it is separated from all other Chlenias spp. listed here, except G.188. (Head
notably less bulbous than in G.188 or G.106, and strongly-marked with dark brown
longitudinal lines on a pale pinkish-tan ground color.) Disturbed fullgrown larvae
are quick to curl inward; if dropping from the plant they fall free, never on a silk
thread. Adults (A) fly March—July (peak June); univoltine. (Preserved = 1-6, 8hh,
Oe Solitons == Il, Sy re, (3)

e Chlenias sp. nov. (det. SF )—G.120. S.AUST., Blackwood (NM): Larvae scarce
(Sept. 65) on Pultenaea largiflorens var. latifolia (only); readily accepted young
Ivs. of Acacia pycnantha as a substitute (dets. NM): These Chlenias larvae show a
confusing mixture of the characteristics of my G.106 and G.103, with respect to
color, maculation, morphology, and behavior. The few adults seen were hard to
separate from G.106, but slightly smaller. Could this be a natural hybrid of
G103 «.G.106P (€Preserved: = 1; 5, 6;°9))

© Chlenias sp. nov. (det. SF)—G.148. S.AUST., +8 mi. NE of Two Wells, at
edges of sandhills (NM & TN): Larvae (20 Aug. 66) abundant on oldman saltbush,
Rhagodia parabolica R. Br—CHENOPODIACEAE (det. MK) which is apparently
the preferred foodplant of this population; also encountered, in far lower numbers
on a few other unrelated native woody plants in this and nearby localities. In gen-
eral appearance, maculation, and behavior, these larvae show strong affinities with
my G.103, but are easily separated and almost certainly another species (or a sub-
species at the least). Two adults emerged: 9 May 67 (4) and 13 May 67 (@);
probably univoltine. (Preserved = 1, 4, 5, 6, 9; photos = 1.)

e Chlenias sp. nov. (det. SF)—G.188 & 188A. (1) W.VIC., 5 mi. S of Kiata,
Lowan Reserve, nr. the designated camping area (NM): Larvae (28 Sept. 67) on
the woody shrub, Baeckea behrii (Schldl.) FvM.—MYRTACEAE (det. MK). These
larvae are very colorful and fairly conspicuous on the foodplants; general appearance
(build), bulbous head, and behavior show affinities with my G.106. All colors and
markings on these larvae are gaudy, rich, and intense—more so than in any of the
other Chlenias larvae here listed (with the possible exceptions of G.111 and G.220,
which look nothing like this one). Two distinctive features: the true legs are vivid
reddish-purple, and there are 6 black dots in a semi-circle directly behind the head
(cervical area). (Preserved = 5, 8h, 9.) (2) S.AUST., 1.2 mi. S of Monarto
South railway crossing, + 10 mi. W of Murray Bridge (NM & TN): Larvae fairly
common (8 Oct. 67) on Baeckea behrii (det. NM). Two é@ adults emerged: 19, 21
May 68; probably univoltine. (Preserved = ¢1, 6, 9; photos = 14.)

e Chlenias sp. (det. NM )—G.206 & 206A. (1) W.AUST., 13 mi. E of Tambellup
(NM & N. B. Tindale): Larvae (21 Nov. 68) on a woody shrub, Melaleuca uncinata
R. Br. ex Ait—MYRTACEAE (det. NM). (Preserved = 5, 6.) (2) W.AUST., at
Wave Rock, nr. Hyden (D. & NM): Larvae (28 Oct. 71) common on a tall
Leptospermum sp. (det. NM); the foodplant was growing abundantly around the
parking area (nr. base of Wave Rock) and was in bloom on this date (petals
pinkish-tinged white). These larvae show affinities with my G.106, and even more

SUPPLEMENT TO VOLUME 33 27

with G.188, but are quite distinct. Pupae were obtained but all died later; no adults
have been seen thus far, but the distinctive larvae are clearly different from all the
foregoing spp. and warrant mention here. Probably univoltine.

General remarks on the foregoing Chlenias spp. (with the exception of G.127 and
G.221, which probably belong in the Oenochrominae): The several incomplete de-
terminations are included here primarily because of the excellent and consistent larval
differences and the diverse (but sometimes specific) foodplant preferences; these
records could be of great immediate assistance to future workers attempting to study
this difficult genus. It took me 5 years and countless field-trips to gather the in-
formation here recorded, all of which is documented by extensive notes, preserved
larvae and pupae, and the associated reared adults; these have been deposited in
the institutions listed in the Introduction. The adults are (in most of the spp.) of
rather similar appearance, somewhat variable, and often tend to overlap in certain
features of the maculation. They are regularly and thoroughly mixed up in most
collections!

Most Chlenias larvae make little or no attempt to hide by day and are not
cryptically-colored—thus usually conspicuous on their foodplants. Good differences
(and affinities) can be seen in the larval head capsules of the various species. Eggs
were not obtained from 5 of the above spp., so little can be said here of their
comparative or taxonomic value. The eggs would certainly warrant close examina-
tion in any future study of this complex and fascinating genus, which is so well-
represented across southern Australia.

@ Ciampa arietaria (Gn.) (det. IC)—G.93. (1) S.AUST., Adelaide city suburbs;
also Blackwood-Belair district (NM): The rather noctuiform larvae (June—Aug.)
are often locally abundant on the weedy South African annual, *Arctotheca calendula
(det. NM). It would be interesting to know what was the primary foodplant here
prior to the introduction of the now exceedingly abundant Cape weed; this moth
may have been considerably less abundant long ago. Adults (A+) fly mid March—
early June; univoltine. The eggs are rain-hatched. See Common (1966b: 81 and
1970: 844) for $ adult photos. (Preserved = 1-6, 9; photos = 1, 2,5.) (2) W.
AUST., Moresby Range, Howatharra Hill Reserve, + 19 mi. NNE of Geraldton (see
Meradend. 1977): One last instar larva (26 Aug. 78) feeding on inflorescence of
*Zaluzianskya divaricata (Thunb. ) Walp.—_SCROPHULARIACEAE (det. NM), at
edge of paddock S of Zone 4 (NM & Lisa Green). This sp. also comes commonly to
uv. light at Drummond Cove, +7 mi. N of Geraldton, W. Aust., (+ mid April-—
July).

e Cleora bitaeniaria (LeGuillou) (det. IC)—G.86. S.AUST., Blackwood (NM):
Captive larvae (April-May) readily accepted mature lvs. of Eucalyptus odorata
(det. NM); larvae are probably present on eucalypts during most months of the
year in this locality. Adults (B+) recorded for every month (peaks Sept., Dec., &
Feb.—June). See pa atend (1972b: 239, 245) for egg photos. (Preserved = 1-6,
Seephores — 1) Ic,-2, 5, 6:)

e@ Cleora displicata Walk. (det. SF)—G.165. (1) S.AUST., Blackwood (NM):
Captive larvae (March) readily accepted young lvs. of Acacia pycnantha (det. NM).
Adults (B) fly Oct—April (peak Jan—March). First instar larval dispersal is notable
in this species. See McFarland (1972b: 239) for egg photos. (Preserved = 1-6, 9;
photos = 1, 2, 6.) (2) S.AUST., 4 & 5 mi. E of Two Wells, at Lewiston Park and
along roadside (NM & TN): Larvae (19 March 67), of what is probably this sp.,
fairly common on Acacia salicina Lindl., A. ligulata A. Cunn. ex Benth. and Cassia
nemophila Cunn. ex Vogel—_MIMOSACEAE & CAESALPINIACEAE (dets. MK).

e “Cleora’ repetita Butl. (det. IC)—-N.QLD., 14 mi. N of Cairns, at Clifton Beach
(D. & NM): Larva (9 May 72) on buds, fls., and lvs. of the shrub, Fenzlia obtusa
Endl.—_MYRTACEAE (det. BH). The adult (a large 2) emerged 19 June 72;
deposited in A.N.I.C., Canberra. (Preserved = 1, 6. )

© Corula seometroides Walk. (det. SF)—G.170 & 170A. CES AUST 2 4eanr
of Two Wells, Lewiston Park (NM & TN; NM & C.N. Smithers): Larvae (19 March

28 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

67: 4 May 67; Sept.—Oct. 69) on native “pine,” Callitris preissii Miq—CUPRES-
SACEAE (det. MK); regularly obtained by beating. Probably multiple-brooded;
some adults emerged Jan._Feb. 1970. (Preserved = 1, 4-6, 9; photos = I, 5, 6.)
(2) VIC., Grampians Mts., S of Hall’s Gap, at Mirrantawa Gap (NM & E. C.
Jaeger): Larvae (26 Sept. 67) on Callitris rhomboidea R. Br. ex Rich. (det. MK).
(Preserved = 5, 8h.)

Criomacha—see Fisera.
e “Diastictis’ goniota Lower (det. NM)—G.214. S.AUST., Hallett Cove, S of
Adelaide (D. & NM): Larvae (2 June 71) abundant on the woody, perennial vine
(not a parasite), Muehlenbeckia gunnii (Hook. f.) Walp.—POLYGONACEAE,
which was growing in tangles over shrubs of Olearia axillaris (DC.) FvM (dets.
NM), just back from the beach. This tentative record has yet to be positively
verified by reared adults, but is probably correct; it is included here because of the
distinctive foodplant which might provide a useful clue toward further investigations
of this moth. The generic placement is doubtful, but the type of goniota is in the
S. Aust. Museum (Adelaide). Adults fly (?)July—Sept.; probably univoltine. (Pre-
served = 1-5, 9; photos = 1.)
e Didymoctenia exsuperata Walk. (det. SF)—G.122. S.AUST., Blackwood (NM):
Captive larvae (Sept.—Oct.) readily accepted young lvs. of Eucalyptus odorata ( det.
NM); larvae are probably present on young lvs. of eucalypts during most months of
the year in this locality. Adults (A) fly Sept._July (peaks Oct-Dec. & May—June)
with by far the majority coming to uv. light only after 2300 hrs. (Preserved = 1-7,
9: photos =. 1) 2.)
e Ectropis excursaria (Gn.) (det. IC, SF )—G.75, 75A, 75B, & 75C. (1) S.AUST.,
N Adelaide city parklands (NM): Larvae (Jan._Feb. 65) common on tender young
sucker-growth at the base of an old pepper tree, *Schinus molle L.—ANACARDI-
ACEAE (det. NM). (2) S.AUST., Belair (F. J. Mitchell): Larvae (Aug.—Oct.)
abundant on young and semi-mature lvs. of the perennial native vine, Hardenbergia
violacea (det. NM); almost defoliating this plant which was being grown against
a garden wall. (3) S.AUST., Blackwood-Belair-Eden Hills district (NM): Larvae
(spring—autumn) on numerous unrelated woody and herbaceous plants (both native
and introduced). Some specific records: In gardens on an ornamental ivy, *Hedera
sp.—ARALIACEAE; *Pelargonium sp.—GERANIACEAE; orange tree (young lvs.
of sucker-growth), *Citrus—RUTACEAE; *Polygonum aviculare L.—POLYGONA-
CEAE; young lvs. of *Eucalyptus cladocalyx FvM.; also (in the natural scrub) often
on Pultenaea largiflorens var. latifolia, new lvs. of Acacia pycnantha, Exocarpos
cupressiformis, Bursaria spinosa, and etc. (dets. NM). Adults (B+) fly all months
(peaks Sept.—Oct., April-June); sexual dimorphism is evident. First instar larval
dispersal is notable in this species. (Preserved = 1, 2, 5, 6, 8h, 9; photos = 1, 2, 5,
6.) (4) S.AUST., 5 mi. E of Two Wells, along roadside (NM & TN): Larvae
(19 March 67) fairly common on Acacia ligulata A. Cunn. ex Benth. and Cassia
nemophila Cunn. ex Vogel—MIMOSACEAE & CAESALPINIACEAE (dets. MK).
(5) W.AUST., Perth, in the California Section of the Botanic Gardens at King’s
Park (D. & NM): Larvae (16 Oct. 72), probably of this sp., feeding (and thriving )
on California sagebrush, *Artemisia californica Less—ASTERACEAE and also on
deerweed, *Lotus scoparius (Nutt. in T. & G.) Ottley—FABACEAE (dets. NM);
both of these plants are abundant natives in the sage scrub and chaparral (evergreen
sclerophyll) associations of southern California, at lower elevations near the coast.
e Ectropis odontocrossa Turner (det. NM)—G.153. S.AUST., + 8 mi. NE of Two
Wells, on white sandhills (NM & TN): Larvae (27 Aug. 66) common on the
woody shrubs, Bertya mitchellii (Sond.) Arg —EUPHORBIACEAE and Dodonaea
bursariifolia FvM. (dets. MK), but not found on any other shrubs beaten in this
habitat. A @ adult emerged 9 June 67. (Preserved = 1, 4-6, 9; photos = 1, 5c.)
© Ectropis pristis Meyr. (det. NM)—W.VIC., 5 mi. S of Kiata, at the Lowan Re-
serve (Sanctuary), nr. the designated camping area (NM): Larvae (28 Sept. 67)

SUPPLEMENT TO VOLUME 33 29

on Baeckea behrii (Schldl.) FvVM.—MYRTACEAE (det. MK). ¢@ and @ adults
emerged, but hatching-date was not recorded. (Preserved = I, 6.)
e Ectropis sp., close to aganopa Meyr. (det. IC)—(1) S.AUST., Yorke Peninsula,
nr. S end of Formby Bay (NM, N. B. Tindale & P. Aitken): Larva (2 Nov. 65) on
Acacia calamifolia Sweet ex Lindl. (det. MK). A @ adult emerged late Nov. 65.
(Preserved = 1, 6.) (2) S.AUST., +4 mi. S of Ashbourne (J. O. Wilson): Larva
(31 March 69) beaten from the twining parasite, Cassytha pubescens (det J. O.
Wilson). An adult emerged 3 May 69; in Wilson Collection, A.N.I.C., Canberra.

Fisera—see also remarks under Mnesampela fucata and Stathmorrhopa macroptila.
e Fisera eribola (Guest) (det. IC)—G.90 (possibly synonymous with Mnesampela
dictyodes Lower). S.AUST., Blackwood (NM): Captive larvae (May-July) readily
accepted mature lvs. of Eucalyptus odorata and E. leucoxylon (dets. NM). Adults
(A) fly late Feb.mid May (peak late March—-early April); univoltine. They come
to uv. light mostly after 2300 hrs. Only minor individual variation is shown in color
and maculation. The eggs seem to be on the borderline of a rain-hatching tendency,
and first instar larval dispersal occurs. The adult rests in the manner of S. macroptila
(p. 32). (Preserved = 1-6, 9; photos = 1, lc, 2, 4, 5, 5c.)
e Fisera perplexata Walk. (det. IC)—G.95 (possibly synonymous with belidearia
Felder). S. AUST., Blackwood (NM): Larvae (July—Oct.) on mature Ivs. of
Eucalyptus odorata (det. NM). Adults (B+) fly April-July. There may possibly
be two distinct spp. involved here, although I suspect it is one moth with highly
variable coloration. Three major color-maculation forms (with minor variations
in each) occur consistently in the Blackwood-Belair district: (a) the earliest form to
appear (April-May) has a pale tan forewing groundcolor with a prominent (darker
brown) single discal dot, a fairly strong, darker brown postmedial line, and little or
no speckling; (b) the next form(s) to appear (from about early May onward) are
generally more-or-less heavily-speckled with darker brown, on medium brown to
orange-brown or gray-brown groundcolor, and the postmedial line is non-existent or
very faint; (c) the last form to appear (from about early June onward) is often a
little larger and very much darker in groundcolor, which can be a deep but dull
reddish- or almost purplish-brown, with little or no evidence of either the post-medial
line or the heavy speckling, although a single discal dot may (or may not) be present.
The flight seasons of forms (a) and (b) overlap (early—late autumn), as do forms
(b) and (c), but (a) and (c) are well-separated. Further rearings, from known
females of all three major forms, all from the same locality, could provide the in-
formation needed for a deeper understanding of this (or these) interesting and per-
plexing species! The eggs are rain-hatched. (Preserved = 1-7, 9; photos = 1 for all
forms, 2 & 5 of earliest form only.)

Gastrinodes—see Cleora.
@ Gastrinopa xylistis Lower (det. NM)—G.137. S.AUST., Blackwood (NM):
Captive larvae (Nov.—Jan.) readily accepted only young lvs. of Eucalyptus odorata
(det. NM). Adults (B+) fly Sept—Feb. (peak Oct—Dec.). First instar larval
dispersal is notable in this species. (Preserved = 1-7, 9; photos = 1, 2, 5, 6.)
e Heteroptila argoplaca (Meyr.) (det. SF, IC)—G.108A. S.AUST., Blackwood
(NM): Captive larvae (most months) readily accepted mature lvs. of Eucalyptus
odorata (det. NM). Adults (A) fly all months (peaks Feb.-March & May-June).
(Preserved = 1-6, 9; photos = 1, 2.)
© Idiodes apicata Gn. (det. SF, IC)—G.147, 147A, & 147B. (1) S.AUST., Black-
wood-Belair district, Highbury (E of Adelaide), and Naracoorte (TN, D. & NM, and
K. J. Sandery): Larvae (most months, esp. winter-spring) on both young and old
(tough) fronds of bracken-fern, Pteridium esculentum (det. Hj. Eichler). This is
apparently the only foodplant of this moth; but see also Metrocampa. Adults (B)
fly all months (peak June—Oct.). See Tillyard (1926: Pl. 39) for adult photo; Mc-
Farland (1972b: 237, 246) for egg photos. (Preserved = 1, 2, 5, 6, 9; photos =
1, Ic, 2.) (2) N.QLD., Atherton Tableland (+ 2500’ el.), nr. Tinaroo (D. & NM):
Larvae (April-May 72) on Pteridium esculentum; also evidence of earlier feeding on

30 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

P. Paquilinum (L). Kuhn (dets. BH) but no larvae found. This latter bracken is
of somewhat taller and more rank growth, and has notably pubescent stems in this
locality. (Both plants are growing together here.) Captive larvae readily accepted
both, but appeared to prefer P. esculentum. (Preserved = 1-6, 9.)
e Lophothalaina habrocosma (Lower) (det. NM )—G.154, 154A, & 154B. S.AUST.,
Mt. Lofty Range, Belair Nat. Park, +1 mi. E of Belair railway station, just S of
Sheoak Rd.; also 4 mi. S of Ashbourne (J. O. Wilson), and nr. Longwood (NM &
TN): Larvae (June-Sept.) on old, mature (sclerophyll) lIvs. of Leptospermum
myrsinoides (det. NM). Adults (D, at Blackwood) recorded only for mid April 67;
probably more abundant at the above-named specific localities, in close association
with the foodplant (a “frail” moth and a weak flyer). Emergences in captivity were
all in March or April (strictly univoltine). The J. O. Wilson Collection, now in
A.N.I.C. (Canberra), contains a good series of reared adults, most of which are
from larvae I collected at the above Belair locality in July—Aug. 68, a winter in
which larvae were very abundant there. The eggs seem to be on the borderline of
a rain-hatching tendency. See McFarland (1972b: 241) for egg photos. See p. 13
regarding adult resting-position. (Preserved = 1-7, 9; photos = 1, 2, 5, 5c, 6.)
e Loweria platydesma (Lower) (det. IC, NM)—G.152. S.AUST., +5 mi. E of
Two Wells (+ 25 mi. N of Adelaide) (NM & TN): Larvae (27 Aug. 66) common,
in a restricted roadside strip, on buds, fls., and mature lvs. of Cassia nemophila Cunn.
ex Vogel—CAESALPINIACEAE (det. MK). Adults emerged mid April 67 (3¢ 6,
12); these 4 specimens show tremendous variation in the darker maculation of the
forewings, but not in the grayish groundcolor. Probably univoltine. (Preserved =
> Clee Os pnotosm— ily)
e@ Melanodes anthracitaria Gn. (det. NM)—G.161. S.AUST., Blackwood-Upper
Sturt district (NM): Captive larvae (Nov.—Dec.) readily accepted only young lvs.
of Eucalyptus odorata (det. NM); no doubt feeds on many other Eucalyptus spp.
as well. Adults (B, at Blackwood) fly mid Sept.—Dec. (peak Nov.); they are more
abundant around Upper Sturt; univoltine. First instar larval dispersal is notable in
this species. See Common (1970: 849, fig. D) for a line drawing of the pupa. See
McFarland (1972b: 241) for egg photos. (Preserved = 1-7, 9; photos = 1, lc, 2, 5,
5c, 6.)
e Metrocampa ada (Butl.) (det. IC)—G.136. S.AUST., Naracoorte, 13 Lochiel
Ave. (TN): Larvae (July—Aug.) common on mature lvs. of bracken, Pteridium
esculentum (det. Hj. Eichler). These larvae are quickly separated from the two
other South Australian bracken-feeding geometrids (Idiodes apicata and Metrocampa
biplaga) by their much smaller size when fullgrown, and by the strong, dark brown,
dorsal chevron markings (= “herring-bone” maculation). Like M. biplaga, the
larvae of ada are notably sluggish in behavior, in contrast to the active larvae of
Idiodes. (Preserved = deformed 1, 5, 6, 8h, 9; photos = 5.)
e Metrocampa biplaga Walk. (det. NM)—G. 217 & 217A—(possibly synonymous
with glaucias Walk.). (1) S.AUST., Naracoorte, 13 Lochiel Ave. (TN): Larvae
(July—Aug.) on older (tough) lvs. of Pteridium esculentum (det. Hj. Eichler).
These larvae, when filled out in last instar, are large and rather “heavy” in build,
and are notably sedentary in behavior; these two characteristics alone will quickly
separate them from larvae of the more common Idiodes apicata (my G.147) which
also occur on the same foodplant, often together with larvae of both Metrocampa
spp. Adults fly March-April (little data available for Naracoorte). (Preserved =
5, 6, 9; photos = 5, 6.) (2) S.AUST., Belair Nat. Park, nr. Long Gully (NM):
Larvae (Sept—Oct. 69) of M. biplaga in company with larvae of I. apicata, both
feeding primarily on the older lvs. of P. esculentum. Idiodes was more abundant at
the time. (Preserved = 1, 2,9; photos = ¢1 from Aldinga, S. Aust.)
Mnesampela—see also Fisera.
e Mnesampela fucata (Felder) (det. NM)—G.109. S.AUST., Blackwood (NM):
Larvae (June—Aug.) on mature (tough) lvs. of Eucalyptus odorata (det. NM);
the unique last instar larva rests outstretched on a strong silken mat, usually on the

SUPPLEMENT TO VOLUME 33 a1

upper (or more exposed) surface of a leaf, always aligned + parallel to the midrib,
in a manner reminiscent of many Papilio larvae. (The same habits are also seen in
the mature larvae of Fisera eribola and F. perplexata, but the perplexata larva is more
often inclined to rest on a leaf margin, especially when smaller.) Adults (A+) fly
mid March-—late June (peak mid April-May), with predominantly ¢ ¢ coming to
uv. light in the earlier part of the flight season and 9 2 mostly appearing from May
onward; ¢ ¢ to light mostly after 2300 hrs, and @@ mostly before that time;
univoltine. The adult forewing shows considerable variation in 2 features: (1)
The groundcolor can vary from pale cream (fresh, unfaded ¢ ¢ only), to light
golden-tan ( ¢ 2), to deep tan with a faint pinkish tinge ( 2 @ only), to rich rust
or orange-brown (@ @ only); (2) the degree of speckling (small darker dots) can
vary from heavy and intense to nil, and is primarily a feature of the 92. The
eggs are rain-hatched and first instar larval dispersal is notable. This species provides
a splendid example to illustrate both of these phenomena, which are described at some
length in an earlier paper (McFarland, 1973). See McFarland (1972b: 239) for
egg photos. (Preserved = 1-7, 9; photos = 1, 2, 5, 5c.)
@ Mnesampela lenaea Meyr. (det. IC)—G.118 & 118A (probably synonymous with
comarcha Meyr.). (1) S.AUST., Blackwood (NM): Larvae (July—Sept.) usually
on saplings (eating only old lvs.) of Eucalyptus odorata or E. leucoxylon. I also
have one record (10 Sept. 66), of a single last instar larva thriving on *E. ficifolia
F. Muell. (W. Aust. native) in a Blackwood garden; this larva had a deep reddish-
pink groundcolor, and was eating only the young (red) lvs. of that highly sclerophyll
eucalypt (dets. NM). These larvae are semi-gregarious when small, but unlike
M. privata have no nest-building habit whatsoever in any of the instars; usually
solitary when fullgrown. They tend to sit exposed on brown or tan (dead) parts of
partially-damaged lvs. in full view, but they are well protected by their close re-
semblance to brown bits of dead and curled leaf. Adults (B-++) fly late March—June
(peak May); univoltine. The eggs are rain-hatched. (Preserved = 1-7, 9; photos =
if ooe G6) 12) SAUST. nur Mt. Lofty Summit (Cleland Park): Larva (19
Aug. 69) on mature lvs. of a Eucalyptus viminalis sapling (det. MK). (3) S.AUST.,
Kangaroo Island, 10 mi. W of Vivonne Bay (NM): Larva (3 Sept. 67) on one of
the mallee eucalypts, E. diversifolia Bonpl. (det. MK).
e Mnesampela privata (Gn.) (det. IC)—G.82. S.AUST., Blackwood (NM): Larvae
(May-July) often conspicuous when defoliating small saplings, especially stunted
or “struggling” individuals (eating old, mature lvs.) of Eucalyptus odorata and
E. leucoxylon (det. NM). The larval habits are unusual in the Geometridae, as they
are semi-gregarious even when fullgrown, more so when smaller; they make tough
silk nests in partially-curled Ivs., in which several rest (half-curled and close together )
_ by day, coming out to feed on nearby foliage after dark. Adults (A) fly late Feb.—
May (peak April); univoltine. See Common (1966b: 83) for a ¢ adult photo;
Common (1970: 849, fig. A) for a line drawing of the final instar larva. (Preserved
—wievaohny O-) photos = 1, 5)\6:)
e Niceteria macrocosma (Lower) (det. NM)—G.200. S.AUST., Blackwood (NM):
Captive larvae (April-June) readily accepted mature lvs. of Eucalyptus odorata ( det.
NM). Adults (B—) fly Feb.—early April, mostly coming to uv. light after 2300 hrs
(even by 0300-0400 hrs they are still coming in); univoltine. This is one of the
largest and most colorful geometrids occuring here. See Common (1966b: 72) for
a good watercolor painting of the adult. See McFarland (1972b: 241) for egg photo.
@eresenved — 1-7 9; photos = I, llc 2; 4, 5; Sc, 6.)
© Osteodes Pfictilaria (Gn.) (det. IC)—G.107. S.AUST., Blackwood (NM): Larvae
(March-July—?) on Dodonaea viscosa Jacq— SAPINDACEAE (det. NM). Adults
(B) fly March-May-—? Probably univoltine. (Preserved = 1, 2, 4-6, 8h, 9; photos =
1, 2, 5.)

Paralaea—see Fisera, Mnesampela, Stathmorrhopa.

Paramelora—see “Diastictis” goniota.

Parosteodes—see Osteodes.

32 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

“Protamelora’” (Aust. Museum, Sydney )—see “Diastictis” goniota.

“Pseudopanthera’—see Lophothalaina.

Rhynchopsota—see “Chlenias” rhyncophora Lower.
e Smyriodes aplectaria Gn., 1857 (det. SF)—G.140 & 140A (possibly synonymous
with notodontaria Walk., 1860). (1) S.AUST., Blackwood-Belair district (NM):
Larvae (Aug.—Oct.) are primarily fl. and fl. bud-feeders (if allowed their preference ),
but will also eat the young lvs. of some of the plants involved. Around Blackwood,
by far the preferred foodplant appears to be Pultenaea largiflorens var. latifolia;
they also readily accepted young lvs. of Acacia pycnantha (dets. NM) in captivity,
but I have no field records of larvae on the latter. In Belair Nat. Park (just S of
Sheoak Rd.), I have found them feeding (in about equal numbers) on 3 unrelated
plants: P. largiflorens latifolia; buds & fls. of the parasitic twiner, Cassytha pubescens;
swelling fl. buds of Leptospermum myrsinoides (dets. NM). Adults (B) fly mid
May-—mid July (peak very short, only late May—early June), and they come to uv.
light primarily after 2300 hrs, which may explain their relative scarcity in collections;
univoltine. The eggs are rain-hatched. (Preserved = 1-7, 9; photos = 1-5, 5c, 6.)
(2) S.AUST., Black Hill, at end of Addison Ave., Athelstone, E of Adelaide (TN):
Several larvae (Aug.—Sept. 67) beaten from Cassytha pubescens which was para-
sitizing the shrubby Casuarina muelleriana (dets. TN); the larvae were definitely
eating the Cassytha, not its host, according to Mr. Newbery. (3) S.AUST., S coast
of Kangaroo Island, + 10 mi. W of Vivonne Bay (NM & G. D. Seton): Larvae (9-16
Oct. 66) on fls. and tender young seed capsules of the spiny-intricate shrub, Daviesia
brevifolia Lindl—FABACEAE (det. MK). (Preserved = 5.)
e Smyriodes serrata (Walk.), 9 May 1857 (det. SF)—G.110 & 110A (possibly
synonymous with carburaria Gn., 31 Dec. 1857). (1) S.AUST., Blackwood (NM):
Larvae (Aug.—Oct.) on young lvs. of Acacia pycnantha (det. NM). Adults (A) fly
late May—early Nov. (peak mid June-early Sept.) almost entirely ¢ ¢ coming to
uv. light in the first few weeks of the season, with 2 2 mostly later. However, the
6 6 continue right through, with both sexes occurring in approximately equal num-
bers (at uv. light) during Sept.; univoltine. (Preserved = 1-6, 9; photos = 1, 2, 4,
4c, 5, 5c, 6.) (2) S.AUST., + 3 mi. S of Monarto South railway station (NM & D.
Bakker): Larva (afternoon of 6 Oct. 69) found crawling among lIvs. on Acacia
calamifolia Sweet ex Lindl. (det. MK). (Preserved = 5.)

Stathmorrhopa—see also Fisera and Mnesampela.
e Stathmorrhopa macroptila (Turner) (det. IC)—G.99. S.AUST., Blackwood
(NM): Larvae (June—Sept.) on young and mature lvs. of Eucalyptus leucoxylon and
E. odorata (dets. NM). Adults (A+) fly April-June (peak late April-May); the
majority come to uv. light after 2300 hrs; univoltine. In fresh specimens, the uni-
formly dark sooty-brown of the forewings, thorax, and head, plus the almost total
lack of maculation (but for an obscure dark discal dot in most males) will serve
to distinguish this sp. from Fisera eribola, the only other geometrid with which it
might be confused in this locality. See p. 13 regarding the adult resting-position. The
eggs are rain-hatched. (Preserved = 1-6, 9; photos = 1, 2, 5, 6.)

Note: Higher up in the Mt. Lofty Range (Aldgate district) occurs what is probably
a distinct but very closely-related species, never seen during six seasons at Blackwood:
This moth flies during late Feb.—April, coming to uv. light mostly after 2300 hrs;
the general appearance is close to S. macroptila, but the major forewing veins are
traced with a soft brick-red, and the brown forewing groundcolor of fresh specimens
is not as dark or sooty as in macroptila.
© Stibaroma melanotoxa Guest (det. IC)—G.81. S.AUST., Blackwood (NM):
Larvae (May-—Sept.) on mature lvs. of Eucalyptus odorata (det. NM); in later instars
they are strictly nocturnal feeders, resting by day outstretched (“catocaline style” )
on bark or on other rough, brown areas of the branches and trunk. Adults (A+)
fly late Feb.early July (peak mid March-early June); the majority come to uv.
light after 2300 hrs; univoltine. There may possibly be 2 spp. involved here but
I suspect this is only one highly variable species with respect to both color and

SUPPLEMENT TO VOLUME 33 20

maculation. First instar larval dispersal is notable. See McFarland (1972b: 237)
for egg photos. (Preserved = 1-6, 9; photos = 1, 2, 5, 6.)
e Stibaroma trigramma Lower (det. IC)—G.104 (synonymous with Cleora dolicho-
ptila Turner (det. NM); type ¢ in Qld. Museum, Brisbane). S.AUST., Blackwood
(NM): Larvae (July—Sept.) on mature lvs. of Eucalyptus odorata (det. NM).
Adults (A) fly mid March-early July (peak mid April-May) with the majority
coming to uv. light after 2300 hrs; univoltine. They are quite variable in forewing
maculation (and to a lesser extent in coloration), but do not approach the great
variability seen in S. melanotoxa. The eggs are rain-hatched, and first instar larval
dispersal is notable. (Preserved = 1-7, 9; photos = 1, 2, 5, 5c, 6.)
e Symmetroctena Peutheta Turner (det. NM)—G.134. S.AUST., SW Yorke Pen.,
S end of Formby Bay, + 5 mi. SW of Carribie Homestead, on beach sandhills (NM,
N. B. Tindale, & P. Aitken): Larvae (2 Noy. 65) common on the small shrubby
root-parasite, Exocarpos syrticolus (FvM. ex Miq.) Stauffer (det. MK); feeding
mostly on young growing tips of stems. Adults emerged late Nov. 65. (Preserved =
maa (6, Sdh, 9: photos = 1.)
e Symmetroctena exprimataria Walk. (det. SF)—G.135 & 135A. S.AUST., upper
Yorke Peninsula, nr. Cunliffe (NM & N. B. Tindale); also, 4 mi. E of Two Wells, at
Lewiston Park (NM, TN, & G. Furness): Larvae (most months) abundant on the
large shrubby root-parasite, Exocarpos aphyllus R. Br. (det. MK), feeding primarily
on floral parts and young tips. Adults fly most months. (Preserved = 1, 4, 5, 6,
Shhh, 9; photos = 1, 5.)
e Symmetroctena exprimataria Walk. (det. IC)—G.233. W.AUST., Drummond
Cove, +7 mi. N of Geraldton (D. & NM): Captive larvae (Jan. 73) readily ac-
cepted floral parts and new growth of Exocarpos sparteus R. Br. (det. WAH). Adults
(B—) are multiple-brooded here. (Preserved = 1-6, 9.)
e Symmetroctena leucoprosopa Turner (det. IC, NM)—S.AUST., + 4 mi. E of Two
Wells, Lewiston Park (NM & TN): Larvae (19 March 67) on the tree, Melaleuca
lanceolata Otto—MYRTACEAE (det. NM). Adults emerged April 67. (Preserved
Bis: 62)

Symmiges—see Smyriodes.
e@ Syneora leucanthes Tumer (det. IC)—G.186. S.AUST., +1 mi. N of Grange,
on the coast W of Adelaide, along Military Rd. (NM): Larvae (8 Aug. 67)
abundant on the large shrub, Melaleuca halmaturorum FvM. ex Mig—MYRTACEAE
(det. MK). Adults emerged Sept.—Oct. 67. (Preserved = 1, 4-6, 9; photos = 1, 5.)
e Syneora sp., close to lygdina Turner (det. NM)—S.AUST., N. Flinders Range,
nr. Arkaroola Homestead, in a dry-rocky gulley (NM): Larvae (Oct. 69) on the
large shrub, Melaleuca glomerata FvM. (det. MK). Adult @ emerged Nov. 69.
ei Preserved = IL, 6.)
e@ Syneora sp., close to mundifera Walk. (det. NM)—S.AUST., S coast of Kangaroo
Is., = 10 mi. W of Vivonne Bay, around borders of a small, freshwater lagoon (NM
& G. D. Seton): Larvae (mid Oct. 66) on the dense shrub, Melaleuca oraria J. M.
Black (det. MK). Adults (6,2) emerged 5, 8 Nov. 66. (Preserved = 1, 6.)
e@ Thalaina angulosa Walk. (det. SF, IC)—G.100 & 100A. (1) S.AUST., Blackwood-
Belair district (NM): Larvae (July—Sept.) regularly on young lIvs. of Acacia
pycnantha; captive larvae readily accepted the eastern Cootamundra wattle, *A.
baileyana FvM. (dets. NM). Adults (A) fly mid March-—mid June (peak April—
early May) with very few @ @ appearing prior to early May; univoltine. This is
the only member of the genus I have taken at Blackwood. The eggs are rain-hatched.
See McFarland (1972b: 241) for egg photos. (Preserved = 1-7, 9; photos = 1,
Ie, 2, 5, 5c, 6.) (2) S.AUST., +3 mi. NE of Woodchester, nr. Hartley (NM):
Larvae (6 July 68) common on Acacia brachybotrya Benth. (det. MK), and on
Cassia sp. CAESALPINIACEAE (det. NM). (Preserved = 5; photos = 5, 5c.) (3)
S.AUST., + 5 mi. E of Two Wells, along roadside (NM & TN): Larvae (27 Aug.
66) common on Cassia nemophila Cunn. ex Vogel (det. NM).

34 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

@ Thalaina clara Walk. (det. NM)—G.128. S.AUST., Naracoorte (eggs through
TN): Captive larvae (May—June 70) readily accepted mature lvs. (only) of Acacia
mearnsii DeWilld. (syn. = A. mollissima Black) (det. NM). Based on the larval
maculation, it is apparent that T. clara larvae (unlike T. angulosa or T. selenoea)
are adapted for living on acacias having finely-bipinnate (“feathery”) lvs., although
they might conceivably accept certain of the other Acacia spp. as “forced” substitutes
in captivity. Adults fly + April-May at Naracoorte (TN records); univoltine. The
eggs are rain-hatched. See Common (1966b: 83) and Tillyard (1926: Pl. 33) for
2 adult photos. (Preserved = 2-7, 9; photos = 2, 5, 6.)

e Thalaina selenoea Dbldy. (det. NM)—G.119 (almost certainly synonymous with
punctilinea Walk.). S.AUST., Naracoorte (eggs through TN): Captive larvae
(June-July 69) readily accepted mature lvs. (phyllodes) of the shrubby *Acacia
iteaphylla FvyM. ex Benth. (det. MK); also casually accepted young lIvs. of A.
pycnantha and A. mearnsii, but much preferred the first species. Adults fly + April—
May at Naracoorte (TN records); univoltine. The eggs are rain-hatched. See
Common (1970: 844) for ¢ adult photo. (Preserved = 1, 2, 5-7, 9; photos = 2, 5,

6.)

e Thalainodes macfarlandi Wilson (det. J. O. Wilson)—G.180. N.TERR., 12 mi.
E of Alice Springs (NM & TN): Captive larvae (June—Sept. 67) readily accepted
mature (tough) phyllodes of Acacia pycnantha (det. NM) as a substitute foodplant at
Blackwood, S.AUST. Adults (B+) fly April-June, probably widespread through
many inland, semi-desert localities between Coober Pedy and Alice Springs; almost
certainly univoltine. The eggs seem to be on the borderline of a rain-hatching
tendency. For the original description and photos of this moth, see Wilson (1972).
(Preserved = 1-6, 9: photos = 1, Ic, 2, 5.)

e Zermizinga indocillisaria Walk. (det. IC, SF)—G.185. S.AUST., Blackwood, and
many other districts around Adelaide (NM): Larvae (winter—spring) are wide-
spread through many natural habitat types here, feeding on a wide range of un-
related plants, mostly shrubs or dwarf shrubs; they are never (or rarely) common
in any given locality, but are regularly encountered almost everywhere during bush-
beating. They look rather like small larvae of Ectropis excursaria at first glance. A
few specific foodplant records: Bertya mitchellii (Sond.) Muell, Arg. (S.AUST.,
+3 mi. S of Monarto South, 6 Oct. 69), and Beyeria leschenaultii var. latifolia
Grining (S.AUST., Hallett Cove, Aug. 67)—both EUPHORBIACEAE (dets. MK);
Cassia nemophila Cunn. ex Vogel—CAESALPINIACEAE (det. NM) (+ 5 mi. W of
Two Wells, S.AUST., 27 Aug. 66); Exocarpos cupressiformis and Pultenaea largi-
florens var. latifolia (dets. NM) (Blackwood-Belair district, S.AUST., Aug.—Sept. ).
Adult ¢ 6 (B+, at Blackwood) fly + May-Sept. (peak June), coming to uv. light
mostly after 2300 hrs; the @ 2 are brachypterous, most peculiar, and unable to fly.
See Common (1966b: 83) for excellent ¢ & @ adult photos. I would expect first
instar larval dispersal in this species. (Preserved = ¢ 2 1, 6; photos = @ I, alive.)

Miscellaneous Unidentified Ennominae

® Genus ? sp. ? (probably undescribed) (det. SF, NM)—G.242. (This sp. may be
fairly close to Chlenias, but it appears to warrant generic separation.) | W.AUST..,
Geraldton district, in Spalding Park at Bluff Point, S side of Chapman R. (D. & NM):
Larvae extremely abundant (July-Aug. 73) on new lvs. of Grevillea pinaster Meisn.
(det. WAH). The situation here was very reminiscent of the periodic extreme
abundance of my G.103 (Chlenias) larvae in Blackwood, S. AUST., where Olearia
ramulosa was nearly stripped of foliage some years in certain localized areas. The
way the colorful G.242 larvae blend with the intricate tangle of linear lvs. on their
foodplant is also parallel to the situation in G.103. However, the prominent pair of
shiny, black, subdorsal chalzae on abdominal segment 2 (A-2), repeated again on
A-8, separate these otherwise very “Chlenias-like’ larvae from all Chlenias spp.

SUPPLEMENT TO VOLUME 33 35

listed earlier; some individuals also have a similar (but smaller) pair of chalzae on
A-3, but in others they are totally missing on that segment. The highly variable
adults fly + May-July; univoltine. (Preserved = 1, 4—6, 9; photos = 1, 5, 5c, 6.)

(B) SuBFAMILY GEOMETRINAE (SYNONYM = HEMITHEINAE )

Aelochroma—see Terpna.
e Agathia sp.? (det. NM)—Gm.421. N.QLD., + 14 mi. N of Cairns, Clifton Beach
(D. & NM): Larva (8 May 72) on semi-mature lvs. (on fast-growing stems) of the
sprawling shrub-vine, Alyxia spicata R. Br.—APOCYNACEAE (det. BH). The single
half-grown larva was killed by a parasite. This tentative larval determination is based
on illustrations and foodplant data from Singh (1953). See Tillyard (1926: Pl. 27)
for A. laetata adult in color. (Preserved = 3rd instar larval head capsule and 8h.)
e Austroterpna idiographa Goldfinch (det. NM)—Gm.412. S.AUST., Highbury
East, at 1136 Lower Northeast Rd. (E of Adelaide) (eggs through K. J. Sandery):
Captive larvae (April-May 70) readily accepted young lvs. of Acacia pycnantha ( det.
NM). Adults (B—) fly March-early May & Sept.—Oct. (records from Sandery Coll.,
Highbury ); extremely rare at Blackwood, where only one worn 2 (early May 65) was
taken in 6 consecutive years. (Preserved = 1-6, 9; photos = 1, 2, 5, 6.)
e Austroterpna Pparatorna (Meyr.) (det. IC)—Gm.405. S.AUST., N. Flinders
Ranges, 21-23 mi. E of Copley (NM, P. & A. Taverna): Larvae (25 Oct. 69) on
young lvs. of Acacia rivalis J. M. Black—MIMOSACEAE (det. S. A. Herbarium ).
Adults emerged mid Nov.—Dec. 69. I suspect that my Gm.412 and Gm.405 are one
and the same species, but they have been recorded separately in my larval collection
and notes. Gm.405 is somewhat less strongly-marked than 412, with a pale gray-frosted
effect over the ¢ forewing. Both Gm.412 and 405 show a considerable size difference
between the sexes, the males usually being notably smaller. (Preserved = 1, 2, 4-7, 9;
photoss— 1s le 5. 5c 6.)
e Chlorocoma Passimilis (Lucas) (det. IC)—Gm.76. S.AUST., Blackwood-Belair
district (NM): Larvae (July—Sept.; also autumn) frequent on young lvs. and growing
tips of Acacia pycnantha (det. NM); also on new growth and fls. of Acacia myrtifolia
(records of J. O. Wilson, Aug. 67, Aldinga, S.Aust.; NM & K. J. Sandery, 4 Sept. 71,
Aldgate, S.Aust.). Adults (A) fly Sept—June (peaks Nov. & Jan.—April) at Black-
wood. See Common (1970: 849, fig. C) for final instar larva of this or a closely-
related sp. (Preserved = 1-6, 8h, 9; photos = 1, Ic, 2, 5, 5c, 6.)
e Chlorocoma cadmaria (Gn.) (det. IC, SF)—Gm.130 & 130A. (1) S.AUST., 3%
mi. SE of Blackwood P.O. (NM): Larvae (Sept.—Oct. 65) on the slender woody
shrub, Leptospermum myrsinoides (det. NM). Adults (C) fly Nov.—March at Black-
wood; rare here, but probably more common near the scattered and localized patches
of their foodplant. (Preserved = 1, 5, 6,9; photos = 1.) (2) S.AUST., S coast of
Kangaroo Island, + 10 mi. W of Vivonne Bay, around edge of a small, freshwater
lagoon (NM & G. D. Seton): Larvae (1 Jan. 66) on Darwinia micropetala (FvM.)
Benth.; also (mid Oct. 66) on Melaleuca oraria J. M. Black and young lvs. of M.
gibbosa Labill.—all MYRTACEAE (dets. MK). See Tillyard (1926: Pl. 27) for adult
in color. (Preserved = 1, 4—6, 9; photos = 1.)
e Chlorocoma sp., close to cadmaria (Gn.) (det. NM)—Gm.41l. VIC., Red Hill
(S of Melboume) (eggs through D. R. Holmes): Captive larvae (Feb.—April 70)
accepted mature lvs. of Leptospermum myrsinoides (det. NM), at Blackwood, S. Aust.
(Preserved = 1-6, 9; photos = 5, 5c.)
e Chlorocoma carenaria Gn. (det. IC)—Gm.407. VIC., Red Hill (S of Melbourne )
(eggs through D. R. Holmes): Captive larvae (April-May) readily accepted young

36 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

lvs. of Acacia pycnantha (det. NM), at Blackwood, S.Aust. Adults emerged mid—
late June. (Preserved = 1-6; photos = 1, 5, 6.)

e Chlorocoma dichloraria Gn. (det. SF)—Gm.401. S.AUST., Naracoorte, at 13
Lochiel Ave. (NM & TN): Captive larvae (Jan. 69) readily accepted young lvs. of
Acacia pycnantha (det. NM), at Blackwood. (Preserved = 1-3, 9; photos = 2.)

e Chlorocoma externa Walk. (det. SF)—Gm.155 (in part; see also the following
Chlorocoma sp.). S.AUST., Blackwood-Belair district (NM): These larvae (most
months, especially winter-spring ) are the commoner of 2 closely-related Chlorocomas
feeding on Exocarpos cupressiformis (det. NM). Adults (B+) fly Sept.-May (peaks
Nov. & April), coming to uv. light esp. after 2300 hrs. This sp. is distinguished by
the presence of small, dark discal dots on all wings, and by the lack of any middorsal,
thoracic-abdominal stripe; the darker spots in the fringe (all wings) are a variable
feature. (Preserved = 1, 4-6, 8h, 9; photos = 1, 2, 5, 6.)

e Chlorocoma sp., close to externa Walk. (det. SF )—Gm.155 (in part; see also the
preceding Chlorocoma). S.AUST., Blackwood-Belair district (NM): Larvae (most
months) on Exocarpos cupressiformis (det. NM). Adults (B—) fly Sept.-May, coming
to uv. light esp. after 2300 hrs. This moth is distinguished from the preceding by the
lack of any dark discal dots, by the presence of a light reddish, middorsal, thoracic-
abdominal stripe, and by a more prominent pale cream border on the outer margins
of both forewings and hindwings, just inside the fringe; darker spots in the fringe
(all wings ) are a variable feature. The two spp. are essentially identical in size (which
varies). The genitalia of this moth and the preceding were found to be distinct (SF
and K. Brookes, Nov. 70). It could be of great interest for a future lepidopterist to
rear several long series of adults from eggs obtained in captivity from known females
of both this form (middorsal stripe) and the preceding (discal dots; no stripe); I had
hoped to do this when living at Blackwood, but never got around to it. (It is possible
that, among my field-collected preserved larvae coded under Gm.155, both forms
may be present, so I would caution anyone who might borrow this material to have
this possibility in mind. It is, however, probable that most or all of the preserved
alcoholic material is referable to the preceding sp., if these two forms are distinct. )
(Preserved = ?, 9; photos = 1, Ic.)

e Chlorocoma m. melocrossa Meyr. (det. SF)—Gm.113. (1) S.AUST., Blackwood-
Belair district (NM): Larvae (Sept.—Oct.) fairly common on the dwarf shrub, Pul-
tenaea largiflorens var. latifolia (det. NM). Adults (B—) fly Oct.-Nov. & mid March—
mid April (peak Oct.). The vivid green adult cannot be mistaken for any other
“emerald” occurring here, if careful note is taken of the head and fringe coloration.
(Preserved = 1, 4-7, 8h, 9; photos = 1, 5, 5c, 6, 6c.) (2) S.AUST., Mt. Lofty Range,
nr. Upper Sturt (TN): Larvae (24 Sept. 67) not uncommon on the dwarf shrub,
Dillwynia hispida (det. TN).

e “Chlorocoma” sp. (det. SF, NM)—Gm.168. (This sp. may warrant generic sepa-
ration from Chlorocoma.) S.AUST., + 5 mi. E of Two Wells, along roadside (NM
& TN): Larvae (Dec.—March) common on Acacia ligulata A. Cunn. ex Benth.; a few
also on a single A. salicina Lindl. (dets. MK). Adults emerged April-May 67. The 46
is the smallest immaculate, soft green geometrid occurring near Adelaide and could be
mistaken for no other; the larval head capsules are also quite distinctive in this sp.
(Preserved = 1, 5, 6, 9; photos = 1.)

e Chlorocoma ?vertumnaria Gn. (det. NM)—Gm.191. S.AUST., +4 mi. S of
Monarto South (NM & TN): Larvae (Sept.—Oct. 67) on Acacia sp. with long, + strap-
like and rather thick phyllodes; stems purplish-tinged. Captive larvae (from eggs of
a confined 2 ) readily accepted young lvs. of A. pycnantha at Blackwood. Adults fly
Oct.—Novy. and possibly other months as well. (Preserved = 1-6, 9; photos = 1, 2,
5, 6.)

© “Crypsiphona” eremnopis Turner (det. NM, IC)—Gm.190. (A synonym is Lopho-
thorax alamphodes Turner, with type ? in A.N.LC.; the type 4@ of C. eremnopis is
also in A.N.I.C. Clearly this moth does not belong in Crypsiphona.) S. AUST., + 10

SUPPLEMENT TO VOLUME 33 Bi

mi. W of Murray Bridge, at 244-3 mi. S of Monarto South railway crossing (NM &
TN): Larvae (8 Oct. 67) fairly common on the low-spreading and dense dwarf shrub,
Dodonaea bursariifolia FvVM.—SAPINDACEAE. This unique habitat, formerly rich in
a great variety of native plant spp. (which are not well represented anywhere else near
Adelaide), is now (1973) totally obliterated; it was on private farmland. There pos-
sibly remains some similar scrub country still intact, a little farther south along the
same road (where “Progress” has not yet blossomed), but I do not recall ever seeing
D. bursariifolia in that area. Adults emerged Novy. 67. (Preserved = 1, 5-7, 8dh, 9;
photos = 1, 5,-6.)

© Crypsiphona ocultaria (Donovan) (det. SF; correct spelling of specific name has
only one “c’)—Gm.84, 84A, & 84B. (1) S.AUST., Blackwood (NM): Larvae (most
months; especially May-June) on either mature or young lvs. of Eucalyptus odorata
(det. NM). Adults (A+) fly Aug.June (peak Nov.—April, especially Feb.). See
Common (1966b: 85) for 2 adult photo; Tillyard (1926: Pl. 39) for ¢. (Preserved
= 1-7, 9; photos = 1, Ic, 2, 5-7.) (2) S.AUST., Coorong district, 2 mi. SSE of Salt
Creek (J. J. H. Szent-Ivany): Larva (14 Nov. 67) on a mallee eucalypt, Eucalyptus
diversifolia Bonpl. (det. MK). (Preserved = 5.)

e@ Cyneoterpna wilsoni Felder (det. SF, IC)—Gm.162. S.AUST., Blackwood (NM):
Captive larvae (Jan.—Feb.) readily accepted young lvs. (only) of Eucalyptus odorata
(det. NM). Adults (B—) fly Aug.—April (peak Dec.—Feb.); the majority come to u.v.
light after 2300 hrs; see McFarland (1972b: 233) for egg photos. @ 9 rarely seen.
(Preserved = 1-7, 9; photos = 1, 2, 5, 6.)

e Eucyclodes buprestaria Gn. (det. NM)—Gm.175, 175A, 175B, & 175C. (1) S.
AUST., Mt. Lofty Range, Upper Sturt district (Ironbank Rd.) to Aldgate (Heather
Rd.) (NM, D. Bakker, TN; K. J. Sandery; A. Smith): Larvae (Mar.—April 67; Aug.—
Sept. 69 & 71) fairly common on young, growing (reddish) tips, fls., fl. buds, and frs.
of the slender-wiry parasitic twiner, Cassytha glabella (det. NM), which was mostly
parasitizing sedges (Cyperaceae) and grasses (Poaceae) here, with a few parasitizing
Exocarpos cupressiformis and Leptospermum myrsinoides. The larvae were found in
all 3 situations, but primarily near the ground, where the supporting hosts of their
foodplant were low, clump-forming perennial sedges and/or native grasses. Adults
(B—) fly spring—early autumn (peaks Nov. & March); extremely rare at Blackwood.
See McFarland (1972b: 233) for egg photos. (Preserved = 1-7, 9; photos = 1, lc,
2a oc, 6, 7.) (2) S.AUST., Belair Nat. Park, *% mi. E of Belair railway station
(NM & D. Bakker): Larva (14 Sept. 69) on Cassytha pubescens which was para-
sitizing Leptospermum myrsinoides (dets. NM); larvae apparently rare here. (3) S.
AUST., Aldinga scrub, +3 mi. S of Port Willunga (P. & K. Sandery): Larva (20
Feb. 72) on fl. buds of Cassytha sp. (det. Sandery); this was probably C. glabella
R. Br. (Preserved = 8d) (4) W.AUST., +16 mi. N of Geraldton, nr. Oakajee
(D. & NM): Half-grown larva (28 July 72) on Cassytha sp. (close to glabella R. Br. )
which was parasitizing shrubby Casuarina campestris Diels. (dets. NM). This moth is
fairly common (B) at Drummond Cove, W.Aust., with by far the majority coming to
u.v. light after 2300 hrs; multiple-brooded here (spring—autumn, with peaks in Sept.—
Oct. and mid April—early May).

e Eucyclodes pieroides Walk. (det. IC) —Gm.420. N.QLD., + 14 mi. N of Cairns,
at Clifton Beach (D. & NM): Half-grown larva (9 May 72) on fls. & fl. buds of the
shrub, Fenzlia obtusa Endl MYRTACEAE (det. BH); also one fresh ¢ adult (same
date) was discovered at rest on one of these bushes (collected). Sexual dimorphism
is striking in this species. Adult 2 emerged 29 June 72. See Common (1970: Pl. 8,
fig. L) for excellent color painting of ¢ adult; Tillyard (1926: Pl. 33, 39) for poor
photos of both sexes. (Preserved = ¢ 92 1, 6, 7, 9; photos = 5.)

e Eucyclodes sp. (det. NM)—Gm.413. N.S.W., Oxford Falls (D. Sands): Larvae
(July—Aug. 70) on lvs. of Dodonaea triquetra Andr. (det. Sands). These larvae are
distinct from either of the foregoing Eucyclodes spp., but show a closer kinship with

38 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

E. pieroides. Unfortunately no adults were successfully reared. (Preserved = pre-
sumed 5; photos = presumed 5.)

e Euloxia fugitivaria (Gn.) (det. NM )—Gm.403 & 403A (possibly synonymous with
isadelpha Turner; type from Waroona, W.AUST.). S.AUST., Highbury East, nr. 1136
Lower Northeast Rd., and Athelstone (Addison Ave.), at base of Black Hill (D. & NM,
P. & K. J. Sandery): Larvae (Oct.—March) on the flimsy but upright dwarf shrub,
Pimelea stricta Meisn.— THYMELAEACEAE (det. MK). Adults (B—) fly late Aug.—
Sept.; unlike most Geometrinae here, this sp. is strictly univoltine. (Preserved = 1-7,
Shh 9= photos) —="1 lle) 25, bc) 6)6e;)

@ Euloxia meracula Turmer (det. NM)—Gm.400. S.AUST., Port Lincoln, SE of town
center (Torrens St.), along beach (TN; D. & NM): Larvae (Sept.—Oct. 68 & 71)
common on the “soft” Ivs. of the dense and compact low shrub, Pimelea serpyllifolia
R. Br—THYMELAEACEAE (det. MK). Adults fly + May-June; strictly univoltine.
(Preserved — 1=6) (Sh; 9) photos = 12) 5.165)

e Gelasma centrophylla (Meyr.) (det. NM)—Gm.406. VIC., Red Hill, S of Mel-
bourne (eggs through D. R. Holmes): Captive larvae (Jan.—Feb. 70) reared at Black-
wood, where they casually accepted Leptospermum myrsinoides (det. NM), but all
died by 2nd—3rd instar; this could have been due to unsuitable foodplant, wrong rear-
ing conditions, and/or disease, which of these was not clear. The record is included
only because it could provide a useful clue; these larvae were growing well, up to a
point! Adults fly summer—? (Preserved = 1-4.)

@ Gelasma rhodocosma Meyr. (det. IC)—Gm.416. N.QLD., Atherton Tableland
(+ 2500’ el.), 12 mi. NE of Atherton, nr. Tinaroo Pines Caravan Park (D. & NM):
Eggs and larvae (April-June 72) on young and semi-mature lvs. of Eucalyptus alba
Reinw. ex Blum and E. Ppolycarpa FvM. (dets. BH). Adults emerged April—May 72.
(Rresenveds— | ap OMOG mo»)

@ Gelasma semicrocea (Walk.) (det. IC, SF )—Gm.89 & 89A. (A synonym is Chloro-
coma ipomopsis Lower, in S.A.M., det. NM). (1) S.AUST., Blackwood (NM):
Larvae (most months) on young lvs. of Eucalyptus odorata (det. NM). Adults (A+)
fly Sept._July (peaks Nov.—Jan., March—April, & June); the majority of individuals
come to uv. light after 2300 hrs. (Preserved = 1-6, 9; photos = 1, lc, 2, 5, 6.) (2)
VIC., Red Hill, S of Melbourne (eggs through D. R. Holmes): Captive larvae (Dec.
69) readily accepted young lvs. of E. odorata at Blackwood, S. Aust. (3). This sp.
also appears to be widespread in the forested SW of W.AUST.

@ Heliomystis electrica Meyr. (det. NM )—Gm.409, 409A, & 409B. (1) S.AUST.,
higher elevations of the Mt. Lofty Range, especially Upper Sturt district to Aldgate
(P. Taverna, A. Smith, & NM): Adults (B—) fly late Oct.-March, with the majority
coming to uv. light only after 2300 hrs and even up to dawn. Never seen at Blackwood
in 6 years. (2) VIC., Red Hill, S of Melbourne (eggs through D. R. Holmes): Cap-
tive larvae (April-May 70; reared at Blackwood) readily accepted luxuriant and soft
young lvs. (from recently-burned trunk resprouts) of (only) Eucalyptus obliqua (det.
MK). Also offered were young, semi-mature, and old lvs. of E. odorata, E. leucoxylon,
and E. fasciculosa; the first two were absolutely refused, but fasciculosa was at first
casually accepted; then they became “sickly” on it and were only saved by switching
them over to obliqua, which was the last eucalypt tried. Upon the latter they soon
regained excellent health, feeding avidly, reaching full size, and eventually forming
large and perfect pupae and adults. It seems probable that the distribution of electrica,
at least in the Mt. Lofty Range, may closely follow that of E. obliqua; this eucalypt
does not grow at elevations as low as Blackwood, although it is well represented in
the higher parts of Belair Nat. Park (several miles to the east ), and thence on to Upper
Sturt, Aldgate, and Mt. Lofty, etc. See Common (1966b: 85) for ¢ adult photo.
(Preserved = 1-7, 9; photos = 1, 2, 5, 5c, 6.) (3) An interesting locality record:
2 4 adults came to uv. light in Oct. 71, nr. Hopetoun, W. AUST. (S of Ravensthorpe).
l’orewings (upperside ) were a deep sooty-brown, with the maculation rather obscured.

© Hypobapta barnardi Goldfinch (det. IC, NM)—Gm.402. N.S.W., Como West,

SUPPLEMENT TO VOLUME 33 39

suburb of Sydney (eggs through L. S. Willan & K. D. Fairey): Captive larvae (Novy.
69) readily accepted young and semi-mature lvs. of Eucalyptus odorata (det. NM),
at Blackwood, S.Aust. Adults emerged Dec. 69. (Preserved = 1-7; photos = 1.)
e Hypobapta eugramma (Lower, 1892) (det. IC)—Gm. 125 & 125A. (It is quite
possible that diffundens Lucas, 1891, will prove to be a synonym; if so, this name
would have priority.) S.AUST., Blackwood (NM): Larvae (late spring—autumn) on
young and semi-mature lvs. of Eucalytpus odorata (det. NM). Adults (B+) fly late
Oct.—early April (peak Jan.—Feb.), especially on hot nights. Sexual dimorphism is
fairly noticeable, primarily in the size difference ( ¢ often much smaller). See Com-
mon (1966b: 85 and 1970: 849, fig. P) for ¢ adult photos; McFariand (1972b: 233,
245) for egg photos. (Preserved = 1-7, 8d, 9; photos = 1, 2, 5, 6.)
e Hypobapta percomptaria (Gn.) (det. IC)—Gm.88 & 88A. S.AUST., Blackwood
(NM): Larvae (spring—autumn) on young and mature lvs. of Eucalyptus odorata and
E. leucoxylon (dets. NM). Adults (B—) fly late Sept.—early May (peak Jan.—April).
An aberrant melanic ¢ of this sp. was taken at uv. light in Belair Nat. Park, 1 April
(ijn. Fisher collector. (Preserved = 1-7, 9; photos = 1, Ic, 2, 3, 5, 5c, 6.)
Lophothorax—see “Crypsiphona” eremnopis.
e Metallochlora militaris (Lucas) (det. IC)—Gm.422. N.QLD., Freshwater (N of
Cairns ), at Limberlost Nursery (D. & NM with L. Abell): Distinctive larva (9 May
72) on fls. & fl. buds of the tree, “star-fruit” or “five-corner” (edible), *Averrhoa
carambola L.—OXALIDACEAE (det. at nursery). Adult @ emerged 1 June 72.
(Preserved = 1, 6, 9.)
e Pingasa chlora (Stoll) (det. I1C)—Gm.417. N.QLD., Atherton Tableland (+ 2500’),
12 mi. NE of Atherton, at Tinaroo Pines Caravan Park (D. & NM): Captive larvae
(May—June 72) accepted young and semi-mature lvs. of the rainforest tree, Euro-
schinus falcata Hook. £—ANACARDIACEAE (det. A. Irvine). Adults emerged early
July 72. (Preserved = 1-7, 9; photos = 5.)
e “Pingasa” emiliaria (Gn.) (det. IC)—Gm.419. (The quotation marks around Pin-
gasa are mine; emiliaria does not appear to belong in this genus.) N.QLD., Ather-
ton Tableland (+ 2500’ el.), nr. Tinaroo Pines Caravan Park (D. & NM): Eggs and
larvae (May-June 72) on young and mature lvs. of Eucalyptus Ppolycarpa FvM. and
E. ?drepanophylla—MYRTACEAE (dets. BH); by far the majority were on small
saplings of the former eucalypt, in a grassy-roadside situation (the saplings were re-
sprouts, in response to cutting back about a year before). The mature larvae have a
surprisingly close (superficial) resemblance to the larvae of Crypsiphona ocultaria.
Adults emerged June—early July 72. (Preserved = 1-7, 8h, 9; photos = 5.)
Prasinocyma—see also Gelasma.
e Prasinocyma ocyptera Meyr. (det. SF )—Gm.169, 169A, 169B, & 169C. (1) S.
AUST., +5 mi. E of Two Wells (NM & TN): Larvae (19 March 67) mostly on a
single tree of Acacia salicina Lindl., and a few on A. ligulata A. Cunn. ex Benth. ( dets.
MK). An adult 2 emerged late March 67. (Preserved = 1, 5, 6; photos = 1.) (2)
S.AUST., + 47 mi. N of Hawker, at Commodore Swamp (NM, A. & P. Taverna):
Adults freshly-emerged (24 Oct. 69), coming to uv. light in a dense thicket of Acacia
victoriae Benth. (det. MK); larvae obtained from a confined @ taken here readily
accepted the soft new growth (lvs., frs., and buds) of this acacia. (Preserved = 1-3,
5; photos = 1, 6.) (3) S.AUST., Flinders Ranges, at foot of Aroona Dam, SSW of
Copley (NM): Larvae (3 Nov. 69) on young lvs. of A. salicina (det. NM). (Pre-
served = 1, 5; photos = 1,5) (4) S.AUST., Leigh Creek (NM & G. Gregory):
Adults (2 Nov. 69) at light; confined for eggs. Resulting larvae reared at Blackwood,
S.Aust., where they readily accepted young lvs. of Acacia pycnantha (det. NM).
Preserved — I. 5, 6: photos = 2.)
e Sterictopsis argyraspis (Lower) (det. IC)—Gm.79. S.AUST., Blackwood (NM):
Larvae (most months) on young and mature lvs. of Eucalyptus odorata (det. NM).

40 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Adults (A) fly Oct.-early May (peaks Nov.—Dec. & March). (Preserved = 1-7, 9;
photos = 1, 2, 4, Ac, 5; 6.)

e Terpna sp. nov. (det. SF, IC )—Gm.156, 156A, & 156B. (Adult appearance is quite
close to mniaria Goldfinch, but it is clearly not that species; for illus. of mniaria, see
Common, 1966b: 85.) S.AUST., coastal bluffs at Hallett Cove, S of Adelaide (D. &
NM, TN, K. Sandery ); also nr. American River and Kingscote, Kangaroo Island (NM):
Larvae (peaks June—Sept. & Jan.—March) on the low and dense shrub, Beyeria leschen-
aultii var. latifolia Grining—EUPHORBIACEAE (det. MK); they match the mature
lvs. of their foodplant to perfection, in respect to both dorsal and ventral color-macula-
tion. Adults (B) fly late Sept—March (peak Oct.-early Nov.). We have seen them
in diurnal flight (1100-1200 hrs +) on several occasions (NM and Sandery); this
was not in response to disturbance. They sometimes rest conspicuously on tops of
bushes (usually the foodplant); they may also rest on the lichen-covered cliffs and
rocks of the Hallett Cove habitat, where they would blend in exceedingly well. (The
2 @ match the pale green lichen patches beautifully, but the ¢ ¢ are a much darker
and richer shade of green. The greens of both sexes change and fade rapidly in the
collection, even when specimens are kept in the dark.) (Preserved = 1-7, 8d, 9;
nos = IL Ie BY 5, He, G, 7.)

e Thalassodes pilaria Gn. (det. SF)—Gm.418. N.QLD., Atherton Tableland (+ 2500’
el.), 12 mi. NE of Atherton, at Tinaroo Pines Caravan Park (D. & NM): Captive larvae
(May-early June 72) readily accepted young and semi-mature lvs. of Eucalyptus
Ppolycarpa FvM. (det. BH). Many adults emerged mid-late June 72. (Preserved =
1-7, 9; photos = 5.)

Miscellaneous Unidentified Geometrinae

® Genus? sp.-—Gm.174. S.AUST., + 4 mi. E of Two Wells, Lewiston Park (D. &
NM; TN); also 1 mi. S of Monarto South railway station (NM & TN): Larvae (May—
Nov.) on native “pine,” Callitris preissii Miq.—CUPRESSACEAE (det. MK); often
obtained by beating but never common. Adults have not been seen in the field, nor
were any ever successfully reared, even after several attempts. This record is included
here in order to draw attention to the existence of this small emerald, in hopes that
some future worker will eventually succeed in rearing it, and thus learn its identity.
(Should this ever occur, I would be most interested to hear about it!) This larva
should not be mistaken for that of my G.170 (see Corula, under Ennominae). The
green larvae of both these spp. are present on the same trees, in the same localities,
at the same seasons of the year. However, those of Gm.174 show one of the typical
geometrine larval body-profiles (and head shapes), and are also typically geometrine
in all details of behavior. Their growth-rate is exceedingly slow; larval mortality is
high in captivity if conditions are not suitable in every detail. (Preserved = 4—6, 8h,
9; photos = 4 or 5.)

@ Genus? sp.? (det. IC, SF, NM)—Gm.404 (fairly close to Chlorocoma in superficial
appearance, but probably belongs in another genus). S.AUST., N. Flinders Ranges,
21-23 mi. E of Copley (NM, P. & A. Taverna): Larvae (25 Oct. 69) on young and
mature lvs. of Acacia rivalis J. M. Black (det. S. A. Herbarium); at Blackwood, S.
Aust., where this rearing was completed, they readily switched over to young lvs. of
A. pycnantha for a substitute foodplant. Many adults emerged Nov.—Dec. 69. The
wing shapes (especially of the ¢ ) are unique among the South Australian emeralds;
this alone will separate them from all other immaculate green geometrines here. (An
adult series, representing this same sp., is in the A.N.I.C., collected by I. F. B. Com-
mon, at 28 mi. W of Madura, W.AUST., 30 April 1968.) (Preserved = 1, 5, 6, 9;
photos ==) 255,163)

(C) SupFAMILY LARENTIINAE (SYNONYM = HypRIOMENINAE)

e Chloroclystis destructata Walk. (det. SF )—G.87 & 87A; G.123 (syn. = catastrep-
tes Meyr.; det. SF). S.AUST., 3 mi. S of Monarto South, 4 mi. SE of Wistow, and in

SUPPLEMENT TO VOLUME 33 4]

the Blackwood-Belair district (NM): Larvae (autumn—winter-spring) on fls. and fl.
buds of various unrelated plants, including goldenrod, *Solidago sp —ASTERACEAE
(in a garden); the native Clematis microphylla DC.—RANUNCULACEAE; Acacia
spp.; Bertya mitchellii (Sond.) Muell. Arg—EUPHORBIACEAE (dets. MK, NM).
Adults (B-++) fly most months (Blackwood). (Preserved = 1, 4-6, 9; photos = only
Clematis in habitat. )
e Chloroclystis sp., either anaspila Meyr. or filata Gn. (det. NM)—G.216. S.AUST.,
Blackwood (NM): Larvae (28 Sept. 69) on Ivs. & buds of Pultenaea largiflorens var.
latifolia (det. NM). Adult emerged 13 Noy. 69. (Preserved = 1, 5, 6.)
© Cidaria uncinata (Gn.) (det. IC)—G.96. S.AUST., Blackwood (NM): Larvae
(autumn—winter-spring) on young lvs. and fl. buds of Hibbertia stricta (det. NM).
Adults (B—) April & late Aug.—Oct. See Common (1966b: 83) for ¢ adult photo.
(Preserved = 1-6, 9; photos = 1, 2, 5.)
© Cidaria ?sp. nov. (det. SF)—G.149. S.AUST., +8 mi. NE of Two Wells, on
white sand hills (NM & TN): Larvae (20, 27 Aug. 66) on fl. buds & fls. of Hibbertia
virgata R. Br. ex DC. (det. MK). Emerged 18, 23 Sept. 66. Genitalia studied indicate
that this slightly larger Cidaria is distinct from wncinata; larval differences lend addi-
tional support to this conclusion. More specimens are needed. (Preserved = 1, 5, 6;
photos = 1, 5c.)
e@ Euchoeca rubropunctaria Dbldy. (det. IC, SF )—G.121 (a synonym is risata Gn.).
S.AUST., Blackwood (NM): Captive larvae (Sept.—Oct.) readily accepted Haloragis
heterophylla (det. NM). I am certain Haloragis is the usual foodplant here. Adults
(A) recorded for all months (peaks Sept. & Nov.); very scarce from May-July.
(Preserved = 1-6, 9; photos = 2.)
e “Euphyia’ actinipha (Lower) (det. IC)—G.98. (The quotation marks around
Euphyia are mine.) S.AUST., Blackwood (NM): Captive larvae (June 65) readily
accepted *Medicago polymorpha var. vulgaris (Benth. ) Shinners—FABACEAE ( det.
NM). Adults (D) fly April-May. (Preserved = 1-6, 9; photos = 1, 5.)
e “Euphyia’ squamulata Warren (det. SF)—G.85. S.AUST., Blackwood (NM):
Captive larvae (May—June) casually accepted fls. and young growth of Olearia ramu-
losa (det. NM). Adults (B+) fly mid Feb.—June; probably univoltine. (Preserved
= 14, P75, 9; photos = 2, 2c.)
e “Euphyia’ sp.? (det. IC, SF, NM)—G.196. S.AUST., + 4 mi. S of Ashbourne,
nr. “Tooperang Scrub” (Mr. & Mrs. J. O. Wilson): One larva (6 Aug. 68) on young
growing tips of Casuarina paludosa var. robusta Macklin. (det. MK). Adult ( 2 )
emerged 20 Sept. 68. Only one adult (a fresh ¢ ) was ever taken in 6 years at Black-
wood, on 26 Oct. 65, at u.v. light; probably univoltine. The latter specimen is deposited
in the B.M.(N.H.). This moth could be mistaken for no other sp.; I could find nothing
to compare with it in any of the major Australian collections visited. The prominent
eyes, particularly in the pupa, are one of its notable features. (Preserved = 4 1, 4, 6,
“19> photos = 61,5, 6.)

Hydrelia—see Euchoeca.
© Microdes orichares Turner (det. NM)—S.AUST., Blackwood (NM): Larva (12
Sept. 65) on lvs. of Olearia ramulosa (det. NM). Adult (2) emerged 12 Oct. 65.
(Preserved = 1.)
e Microdes squamulata (Gn.) (det. IC, SF)—G.102; ?G.117. S.AUST., Blackwood
(NM): Larvae (winter-early spring) on young lvs., fls., and buds of several Acacia
spp.; in a garden common every winter on tender young lvs. of A. baileyana FvM.
(det. NM). Adults (A) fly Oct.July (peaks Nov. & May-June). (Preserved = 1,
oo: photos = 1) 5.)
e Poecilasthena ?ischnophrica Turner (det. NM )—G.173 (closely matches a speci-
men labelled “type” in Qld. Museum, Brisbane). S.AUST., Belair Nat. Park, 1 mi. E
of Belair railway station (NM & D. Bakker): Larvae (mid Sept. 69) abundant, feed-
ing on tough, mature lvs. of Leptospermum myrsinoides (det. NM). Adults never
taken at Blackwood; probably restricted to limited areas where the foodplant grows,

42 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

as it is a weak flier. I had never before encountered these larvae on this plant, although
I had been collecting in the same locality every winter-spring (and frequently beating
Leptospermum) since 1965; probably univoltine and cyclic. (Preserved = 1, 5, 6.)
e@ Poecilasthena pulchraria Dbldy. (det. IC, SF)—G.144. S.AUST., Blackwood-
Belair district (NM): Captive larvae (June-early Aug.) readily accepted mature lvs.,
fls., and green fruits of the semi-woody, mat-forming “native cranberry,” Astroloma
humifusum (det. MK). From a vast array of native plants and weeds offered (to two
different batches of these larvae), only A. humifusum proved acceptable, and it was
taken avidly; rapid and healthy growth resulted. This is very probably a correct inter-
pretation of the “preferred” or “true” foodplant of this moth, in the locality named.
Adults (C, at Blackwood) recorded for most months (peak Nov.—Dec.). (Preserved
== 1=6> 9s photoss— a6)

e Xanthorhoe sodaliata Walk. (det. SF)—G.139. S.AUST., Glenelg (J. O. Wilson):
Captive larvae (Sept. 71) readily accepted lvs., salmon-pink fls., and buds of scarlet
pimpernel, a small annual weed, *Anagallis arvensis L.—PRIMULACEAE (det. NM).
I had previously attempted to rear this sp. at Blackwood without success, having
offered the captive larvae various low-weedy, herbaceous plants, but Anagallis was not
among them; other plants, such as clovers, Plantago, etc., were refused. The larvae
reared by Wilson (on Anagallis) were clearly on a preferred foodplant and were
thriving in last instar when I saw them. Adults (B+, at Blackwood) fly Sept.—Dec.;
univoltine. (Preserved = 1.)

e Xanthorhoe subidaria Gn. (det. SF, IC)—G.138. S.AUST., Blackwood (NM):
Captive larvae (July—Aug.) readily accepted a bur clover, *Medicago polymorpha var.
vulgaris ( Benth.) Shinners (= M. denticulata) —FABACEAE (det. NM). Adults (A)
recorded for all months (peak Sept.—_Nov.). (Preserved = 1-6, 9; photos = 5c. )

e Xanthorhoe vacuaria Gn. (det. SF)—G.213 (a synonym = solutata Walk.). S.
AUST., Blackwood (NM): Captive larvae (April-May 69) readily accepted *Medi-
cago polymorpha var. vulgaris (Benth. ) Shinners—FABACEAE (det. NM). I have
no records of adult abundance or flight season for this species. (Preserved = 1-5, 9;
Photos — 2)

@ Xanthorhoe vicissata (Gn.) (det. IC, SF)—G.92. S.AUST., Blackwood (NM):
Captive larvae (May—June) accepted a wide range of unrelated herbaceous plants
(and also one semi-woody dwarf-shrub ), as follows: *Centaurium sp.—GENTIANA-
CEAE; *Chenopodium sp.—CHENOPODIACEAE; Hibbertia sp.; Lythrum hyssopi-
folia L.—LYTHRACEAE; a weedy Malva sp—MALVACEAE; *Medicago poly-
morpha var. vulgaris ( Benth.) Shinners—FABACEAE; *Mentha sp.m—LAMIACEAE;
* Plantago lanceolata L.—PLANTAGINACEAE; *Polygonum sp.—POLYGONACEAE;
fls. of goldenrod, *Solidago sp—ASTERACEAE; and chickweed, *Stellaria sp.—
CARYOPHYLLACEAE (dets. NM); after Ist instar, they were reared to maturity on
a combination of Medicago and Plantago lvs. (both common lawn weeds here) which
they ate avidly. Adults (A) fly March—April (peak late March); univoltine. The eggs
seem to be on the borderline of a rain-hatching tendency. (Preserved = 1-6, 9; photos
== Melle? 5):

(D) SuBFAMILY OENOCHROMINAE (OENOCHROMATINAE )

e@ Arhodia lasiocamparia Gn. (det. IC, SF)—G.124. S.AUST., Blackwood (NM):
Captive larvae (Nov.—Dec.) readily accepted young lvs. of Eucalyptus odorata ( det.
NM); I also have 3 field-records of larvae collected on 3 different (unidentified )
Eucalyptus spp., in the Blackwood-Belair district and at Black Hill, E of Adelaide
(NM & TN). Adults (B+) fly Sept.-mid March (peak late Oct.—Dec.); the majority
come to u.v. light only after 2300 hrs (2 2 rarely seen); univoltine. Color forms vary
considerably; a pale pinkish-tinged form (rarely seen at Blackwood) was often taken
at Upper Sturt. First instar larval dispersal is notable in this species. See McFarland
(1972b: 233) for egg photos. (Preserved = 1-6, 9; photos = 1, Ic, 2, 5, 5c, 6.)

SUPPLEMENT TO VOLUME 33 43

e Arhodia Pretractaria Walker (det. SF)—G.163 (clearly distinct from lasiocam-
paria). S.AUST., + 5 mi. W of Springton (NE of Adelaide) (TN): A single, long,
encircling-band type of egg mass was found attached to a thin stem of a Casuarina sp.
at roadside (late Dec. 66). Resulting larvae (Jan.) reared at Blackwood; they ab-
solutely refused Casuarina spp. offered, but readily accepted young lvs. of Eucalyptus
odorata (det. NM). Adults (3 6 6, 12) emerged late Nov.—early Dec. 68, after 22
months in the pupal stage. (I did not attempt to break the diapause between Sept.—
Dec. 67, as I was away at the time; the pupae thus passed through a second summer
and winter in diapause. Moisture applied from Sept.—Nov. 68 brought out the adults. )
First instar larval dispersal was observed in this species. Probably tnivoltine. (Pre-
served = 1-6, 9; photos = I, 5, 6.)

Cycloprorodes—see “Chlenias” melanoxysta Meyr. (Ennominae ).

Descoreba—see Arhodia.

@ Dichromodes anelictis Meyr. (det. NM )—W.VIC., Lowan Reserve, +5 mi. S of
Kiata (NM): Larvae (28 Sept. 67) on Baeckea behrii (Schldl.) FvM.—MYRTA-
CEAE (det. MK). (Preserved = 41, 6.)

e@ Dichromodes atrosignata Walk. (det. NM)—G.187. W.VIC., Lowan Reserve,
+5 mi. S of Kiata (NM): Larvae (28 Sept. 67) very abundant on Baeckea behiii
(det. MK). Adults emerged Oct.—Nov.; highly variable in coloring and maculation.
(Preserved = 1, 4-6.)

e@ Dichromodes sp., close to eusurpatrix Prout & exsignata Walk. (det. SF, NM)—
G.164. (1) S.AUST., 3%4 mi. SE of Blackwood P.O. and Belair Nat. Park, 1 mi. E of
Belair railway station (NM & TN): Larvae (Oct.—Dec. 66) abundant in the first
locality, on tough, mature lvs. of Leptospermum myrsinoides (det. NM); scarce in the
second locality although the foodplant was common there. Adults emerged early Oct.
67, after 9 months in pupal diapause; univoltine. (Preserved = 1, 4-7, 9; photos =
1, 4-6.) (2) S.AUST., + 1 mi. N of Coonalpyn (NM): Larvae (28 Dec. 68) on
Leptospermum coriaceum (FvM. ex Miq.) Cheel; also present on Leptospermum sp.
at + 4 mi. E of Lucindale, S.AUST., 26 Dec. 68 (NM & TN).

@ Dichromodes explanata Walk. (det. NM)—G.150 & 150A. (1) S.AUST., + 8 mi.
NE of Two Wells, on white sandhills (NM & TN): Larvae (20, 27 Aug. 66) on young
lvs. of the erect shrub, Calytrix involucrata J. M. Black (det. MK). Adults emerged
mid Oct. 66; little variation was evident in the specimens seen. (Preserved = 1,
4-6, 9: photos = 1, 5, 5c.) (2) S.AUST., Athelstone, at foot of Black Hill
(end of Addison Ave.) (NM): Larvae (15 June 68), mostly half-grown, abundant
on Calytrix tetragona (det. NM). (3) S.AUST., S coast of Kangaroo Island,
+ 10 mi. W of Vivonne Bay (NM & G. D. Seton): Larva (12 Oct. 66) on the dwarf
shrub, Lhotskya sp. (= Calytrix) -MYRTACEAE (det. MK). (Preserved = 5.)

@ Dichromodes limosa Turner (det. IC )—G.227. N.QLD., Clifton Beach, + 14 mi.
N of Cairns (D. & NM): Larvae (9 May 72) abundant on fls., buds, & young lvs. of
the shrub, Fenzlia obtusa Endl—MYRTACEAE (det. BH). Adults emerged early—
mim juUne 2. (Preserved = 1, 5, 6, Sh, 9:)

e@ Dichromodes partitaria Walk. (det. NM)—G.158. S.AUST., S coast of Kangaroo
Is., = 10 mi. W of Vivonne Bay (NM & G. D. Seton): Larvae (mid Oct. 66) pri-
marily on lvs. of Melaleuca oraria J. M. Black; a few also beaten from M. gibbosa
Labill MYRTACEAE (dets. MK), but the former appears to be preferred here.
(Preserved = 1, 4-6, 9.)

A comment on adults of the genus Dichromodes: These geometrids appear to
occupy much the same ecological niches in Australia as do adults of Semiothisa
spp. (Ennominae) in North America. They are highly variable as to coloration,
but grays, tans, browns, and red-browns predominate, with nearly all spp. being
cryptically colored and marked to match the soil, rock surfaces, leaf litter, or small
dead leaves, etc. Although essentially nocturnal in activity, they are easily alarmed to
flight in the daytime; the characteristic behavior is to fly up before the walker, only

44 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

to land again on the ground a short distance away, with wings pressed flat against
the ground and the hindwings totally covering the forewings (planniform position ).
Dinophalus—see also Lissomma and “Miscellaneous Unidentified Oenochrominae”
(end of this section).
@ Dinophalus drakei (Prout) (det. IC, SF)—G.194. S.AUST., Athelstone, nr. foot
of Black Hill (end of Addison Ave., E of Adelaide) (TN): The original 2 adult
(15 Oct. 67) came to uv. light; it was given to me and eggs were obtained. Captive
larvae ( Nov. 67) readily accepted only the tender young linear lvs. of the dense, tough-
sclerophyll shrub, Hakea rostrata (det. NM). Adults emerged late Oct. 68; univoltine.
See McFarland (1972b: 235) for egg photo. (Preserved = 1-6, 9; photos = 1, 2, 6.)
e Dinophalus serpentaria (Gn.) (det. IC, NM)—G.193 & 193A (a synonym = un-
dulifera Walk.). (1) S.AUST., 3.3 mi. S of Monarto South railway station (NM &
TN): The original 9 adult (8 Oct. 67) came to uv. light and was confined for eggs.
The resulting larvae were reared at Blackwood, where they readily accepted tender
young lvs. (only) of Hakea rostrata (det. NM); univoltine. (Preserved = 1, 2, 5, 9;
photos = 1, 2,5.) (2) W.AUST., nr. Red Bluff, + 3 mi. S of Kalbarri (D. & NM):
One larva, identical to the above (late Nov.—early Dec. 71), on tender young lvs. of
Grevillea ?trachytheca FvM. (det. NM). A deformed adult ¢ emerged 13 Jan. 72.
(Preserved = 1, 6; photos = 5.)
e Hypographa aristarcha Prout (det. IC, SF)—G.184. (1) S.AUST., S coast of
Kangaroo Island, + 10 mi. W of Vivonne Bay (NM & E. C. Jaeger): The original ?
adult (6 Sept. 67) came to uv. light and was confined for eggs. The resulting larvae
were reared at Blackwood, where they readily accepted fls. and young lvs. (only) of
Hakea rostrata (det. NM); they also accepted tender young lvs. of the highly-sclero-
phyll and spiny low shrub, Isopogon ceratophyllus, but the rearing was completed
using only Hakea, upon which they thrived. Adults of this most peculiar, large geo-
metrid (with a veliform resting position) fly from late winter-spring; univoltine. See
McFarland (1972b: 235) for egg photos. (Preserved = 1-6, 9; photos = 1, 2, 4-6.)
(2) I know of two recent mainland S$. AUST. records as follows: (a) Athelstone, at
39 Addison Ave., nr. foot of Black Hill, 13 Oct. 67, a fresh adult ? at uv. light (J. J.
H. Szent-Ivany); (b) ar. Kingston, 29 Aug. 72, a fresh 2 at uv. light (TN). (3)
I collected 13 mostly fresh adults (12 ¢ 4, 19) of what appears to be a very closely-
related sp. (if it is not, in fact, a ssp. of aristarcha), in W.AUST., in the Stirling
Ranges, nr. Toolbrunup Peak, 19 Nov. 68, at uv. light. (Preserved = 1; photos = 1.)
See p. 13 regarding the adult resting-position.
e “Hypographa” epiodes Turner (det. NM)—G.205. (1) W.AUST., Stirling Ranges,
nr. Toolbrunup Peak, in a roadside gravel-quarry (NM & N. B. Tindale): Larvae (19
Nov. 68) feeding at night on young lvs. of Banksia sphaerocarpa R. Br. (det. WAH).
An adult ¢ emerged 13 Oct. 69, after over 10 months in pupal diapause. (Preserved
= 1,5, 6; photos = 1, lc.) (2) W.AUST., Moresby Range, Howatharra Hill Reserve,
+19 mi. NNE of Geraldton (NM): Larvae (Sept.—Oct.) on young lvs. (only) of
Dryandra fraseri R. Br. (det. WAH).
e Hypographa hiracopsis Meyr. (det. SF)—G.240. | W.AUST., Geraldton-Drummond
Cove district (NM): Captive larvae (July—early Aug.) readily accepted the youngest
new leaves (only) of Grevillea pinaster Meisn.— PROTEACEAE (det. NM); nocturnal
feeders. Adults (B—) fly mid May-—late June; univoltine. (Preserved = 1-6, 9; photos
an)
© Lissomma ampycteria Turner (det. IC)—G.223. W.AUST., + 2% mi. S of Kal-
barri, at Red Bluff Caravan Park (D. & NM): Larvae (mid-late Nov. 71) nocturnal
feeders on tender young lvs. of the common “smoke bush,” Conospermum stoechadis
Endl.—PROTEACEAE (det. NM). An adult ¢ emerged Jan. 72. (Preserved = 1, 5,
6, 8d; photos = 5, 5c.)
© Macrotenia ochripennata ( Walk.) (det. IC )—G.222. W.AUST., on the south coast
at East Mt. Barren, + 6 mi. W of Hopetoun (D. & NM): The original 9 adult (23
Oct. 71) came to uv. light and was confined for eggs. The resulting larvae were reared
at Red Bluff, nearly 700 mi. to the NW of the collection-site, where they accepted

SUPPLEMENT TO VOLUME 33 45

young lvs. of Conospermum stoechadis End|—PROTEACEAE (det. NM). These
larvae appear to have a fairly close kinship with Oenochroma spp. Adults fly Sept.—
Nov.—?; probably univoltine. (Preserved = 1-5; photos = 4.)
e@ Mictodoca Ptoxeuta Meyr. (det. IC)—-W.VIC., Lowan Reserve, + 5 mi. S of Kiata
(NM): Larvae (28 Sept. 67) on Baeckea behrii (Schldl.) FvM.—MYRTACEAE
(det. MK). Adults (1 ¢, 322) emerged early-mid April 68; probably univoltine.
( Preserved = 1, 6.)
e Monoctenia falernaria Gn. (det. IC, SF)—G.167. S.AUST., Blackwood (NM):
Captive larvae (May-Sept.) readily accepted old and semi-mature lvs. of Eucalyptus
odorata (det. NM). Adults (B) fly March—April (peak mid-late March); univoltine.
They come to uv. light almost without exception after 2300 hrs; 2 2 hardly ever seen
at lights. There is considerable variation in groundcolor and forewing maculation; the
usual form here is predominantly light brown or tan, not pink-tinged. A photograph
of a living adult ¢ of this sp., in its usual resting position, appears on the cover of
J. Res. Lepid. 12(4), with data inside back cover. This moth and Phallaria ophiusaria
(p. 46) are the two largest geometrids at Blackwood; a close third and fourth would
be Niceteria (Ennominae) and M. smerintharia respectively. See McFarland (1972b:
235) for egg photos. (Preserved = 1-6, 9; photos = 1, Ic, 2, 4-6.)
@ Monoctenia smerintharia Felder (det. IC, SF)—G.94 (a synonym is probably
calladelpha Lower; type in S.A. Museum). S.AUST., Blackwood (NM): Captive
larvae (May-—Sept.) readily accepted mature (old) lvs. of Eucalyptus leucoxylon and
E. odorata (dets. NM); reared on the latter. Adults (A) fly mid Feb—early May
(peak late March-early April); univoltine. The majority of 2 2 (rarely seen at light)
come in before 2300 hrs; most of the ¢ ¢ come in after 2300 hrs and are abundant
at uv. light here. Differences in the ¢ & 2 forewing coloration and maculation are
notable in this sp.; most ¢ ¢ here are almost immaculate on the forewings and thorax.
See McFarland (1972b: 235) for egg photos. (Preserved = 1-6, 9; photos = 1, élec,
252 oe, 6.)

Nigasa—see Arhodia.

Oenochroma—see also Macrotenia.
® Oenochroma vinaria Gn. (det. IC, SF )—G.77, 77A, 77B, & 77C. (1) S.AUST..,
Blackwood-Belair district and Adelaide city and other suburbs (NM): Larvae (win-
ter-early summer) on various eastern *Grevillea spp. used commonly in gardens as
ornamental shrubs and hedges; among these is also included the tree, silky-oak, *G.
robusta A. Cunn—PROTEACEAE (det. NM). The “wild” populations in native
scrub areas of the Belair district (as opposed to those in the city and suburban gar-
dens), feed almost exclusively on tough-sclerophyll Hakea spp. (needle-bushes ), esp.
H. rostrata (det. NM), eating both old and young lvs., but prefering the latter when
available. (I have never found them on the only native Grevillea of the Belair district,
G. lavandulacea.) The older larvae often rest outstretched (lasiocampid or catocaline-
style) along the lower woody or brown parts of stems, coming up to feed after dark.
Adults (B+) recorded for all months (peaks Oct—Dec., March—May, & late July);
the majority (both sexes) come to uv. light after 2300 hrs. Both sexes are highly
variable in ground color; less so in maculation. A washed-out print showing the usual
resting position of an adult ¢ of this sp. as viewed from two different angles appears
on the cover of J. Res. Lepid. 14(1), with comments on p. 60. See also Common
(1966b: 81) and Tillyard (1926: Pl. 39) for ¢ adult photos; Common (1970: 849,
fig. B) for illus. of final instar larva. (Preserved = 1-6, 9; photos = 1, lc, 2, 5, 6.)
(2) S.AUST., + 8 mi. NE of Two Wells, on whitish sandhills (NM & TN): Larvae
(20 Aug. 66) on Grevillea ilicifolia R. Br. (det. MK). (3) S.AUST., south coast of
Kangaroo Is., + 10 mi. W of Vivonne Bay (NM & G. D. Seton): Larva (10 Oct. 66)
on the very tough old lvs. of Hakea muelleriana Black (det. MK); no new growth
was present anywhere on the plant. (4) W.AUST., on the west coast, 21 mi. S of
Lancelin (NM & N. B. Tindale): Larvae (9 Nov. 68) common, feeding on young
lvs. and fls. of the common and widespread Hakea trifurcata (Sm.) R. Br. (det.

46 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

WAH). (5) W.AUST., + 2 mi. NNE of Howatharra Hill Reserve, or + 21 mi. NNE
of Geraldton (D. & NM): Mature larvae (6 Oct. 74) on the fully-open fls. of Hakea
auriculata Meisn. (det. WAH), very plump and feeding avidly; a few were also eating
tender new lvs. where present, but the fls. were clearly preferred. (6) W.AUST.,
Moresby Ranges, Oakajee district, + 16 mi. NNE of Geraldton (NM): Larvae (13
Oct. 74) on tender new lIvs. of Hakea lissocarpha R. Br. (det. WAH).
e Onychopsis lutosaria (Felder) (det. NM)—G.219. VIC., Red Hill, south of Mel-
bourne (eggs through D. R. Holmes; orig. 9 14 Feb. 70): Captive larvae (March-—
April 70) readily accepted mature lvs. of Eucalyptus odorata (det. NM) at Blackwood,
S. Aust. To my knowledge, this sp. has never been collected in the Mt. Lofty Range.
Sexual dimorphism is notable; a good color illustration of the 2 is given by Common
(1966b: 72). (Preserved = 1-6, 9; photos = 691, élc 5, 66.)
Ophiographa—see Dinophalus and Lissomma.
@ Parepisparis sp. (det. IC)—G.226. N.QLD., Atherton Tableland (+ 2500’ el.),
at Tinaroo Pines Caravan Park (NM & N. B. Tindale): The original 9 adult (10
May 72) came to uv. light; eggs obtained. Captive larvae (late May—June 72) readily
accepted young and semi-mature lvs. of Eucalyptus ?polycarpa FvM. (det. BH). Un-
fortunately, this rearing could not be completed, as we were leaving the locality in
late June; the larvae refused other offered eucalypt lvs. that were encountered in
transit, thus dying. (Preserved = 1-5.)
e Phallaria ophiusaria Gn. (det. IC, SF)—G.80. S.AUST., Blackwood-Belair district
(NM): Captive larvae (June—Oct.) accepted mature lvs. of Dodonaea viscosa; I was
told (by an eastern correspondent ) that this was the only foodplant of P. ophiusaria,
and so did not bother to offer them any of the other local native plants. They
grew fairly well on the Dodonaea, but I have no evidence that they prefer it (or are
ever to be found on it) in this habitat. On several subsequent occasions, however,
larvae were collected at night (Sept., at Blackwood), feeding on tough mature lvs. of
Acacia pycnantha, and during diurnal beating (July—Sept., at Belair) on tough mature
lvs. (no new growth present on the shrubs) of Hakea rostrata (dets. NM). Relatively
few of the “Macro” spp. normally feeding on A. pycnantha or H. rostrata will eat the
very tough oldest lvs., a point of interest in the case of this moth. New lvs., in fact,
do not normally begin to appear on A. pycnantha or H. rostrata until early spring
(Sept-Oct. +), by which time the Phallaria larvae have already been feeding and
growing (very slowly) for 4-5 months. They are mostly in last instar by Sept.—Oct.
in this locality. Adults (B) fly mid Feb.—late March (short peak in early-mid March
only); univoltine. By far the majority of individuals come to uv. light after 2300 hrs;
both sexes are about equally represented at light; if anything, 2 2 come in more often
than ¢ ¢ (the reverse of the usual situation). A moderate degree of variation is seen
in the wing groundcolors (from ashy-gray to brown to pale tan). See McFarland
(1972b: 237) for egg photos. (Preserved = 1—6, 9: photos = 1, 2, 5, be.)
e Phrixocomes sp., nr. ptilomacra Lower (det. NM)—G.157. S.AUST., south coast
of Kangaroo Is., +10 mi. W of Vivonne Bay (NM & G. D. Seton): Larvae (19
Oct. 66) on Melaleuca oraria J. M. Black and M. gibbosa Labill—MYRTACEAE
(dets. MK); the former appears to be preferred in this locality. An adult ¢ emerged
A Dec: 66. (Preserved) = 61) 749756) photoss—= oul)
Rhynchopsota—see “Chlenias” rhyncophora Lower (Ennominae ).

Miscellaneous Unidentified Oenochrominae

@ ?Dinophalus sp. (det. NM)—G.224. W.AUST., on the west coast + 3 mi. S of
Kalbarri, nr. Red Bluff Caravan Park (D. & NM): Larvae (late Nov.-mid Dec. 71)
feeding both day and night on young lvs. of Grevillea Ptrachytheca (det. NM), and
resting among the lvs. close to areas of feeding. No adults were successfully reared.
(Preserved = 5, 6, 9; photos = 5, )

SUPPLEMENT TO VOLUME 33 AT

(E) SUBFAMILY STERRHINAE (SYN. = ACIDALIINAE )

e Chrysocraspeda cruoraria (Warren) (det. IC)—G.228. N.QLD., Atherton Table-
land (+ 2500’ el.), 12 mi. NE of Atherton, nr. Tinaroo Pines Caravan Park. (D. &
NM): Captive larvae (June 72) readily accepted young and semi-mature lvs. of
Eucalyptus ?polycarpa FvM. (det. BH). Completion of this rearing was not possible,
as it was necessary to leave the locality in late June; the larvae died in transit. They
were, however, thriving on the above eucalypt prior to departure. See Tillyard (1926:
Pl. 39) for a ¢ adult photo of the closely-related C. aurimargo Warr. (Preserved =
i429) )

e Eois (Idaea)costaria (Walk.) (det. SF)—G.83; ?G.209. S.AUST., Blackwood
(NM): Captive larvae (May-June) readily accepted buds and fls. of wireweed,
*Polygonum Paviculare L.—POLYGONACEAE (det. NM); the lvs. were also accepted
after about 3rd instar. Adults (A) fly + Oct.-May (peak Nov.—April). (Preserved
=O. 9 photos = 2,5.)

e Eois (Idaea) philocosma (Meyr.) (det. SF)—G.197. S.AUST., Blackwood (NM):
Captive larvae (Jan.—March 69) readily accepted lvs. and floral parts of *Polygonum
Paviculare L. (det. NM). Adults (A) fly spring—autumn (peak Jan.). (Preserved =
t=65.9: photos = 1, 2, 5, 6.)

e Scopula (Acidalia) optivata (Walk.) (det. SF)—G.198. S.AUST., Blackwood
(NM): Captive larvae (Jan.—Feb. 69) readily accepted lvs. of *Polygonum Paviculare
L. and bur clover, *Medicago polymorpha var. vulgaris ( Benth.) Shinners (dets. NM);
of the two, the former was preferred. Adults (A) fly spring—autumn (peak Jan.).
(Preserved = 1-6, 9: photos = 1, 2.)

e Scopula (Acidalia) rubraria (Dbldy.) (det. IC)—G.97. S.AUST., Blackwood
(NM): Captive larvae (May-June) avidly accepted lvs. of the common lawn weed,
plantain or ribwort, *Plantago lanceolata—PLANTAGINACEAE (det. NM). Adults
(A+) recorded for every month (peak Dec.—March); they rest by day among weeds
and grasses, hanging from stems, etc., and are particularly common around weedy,
uncut lawns. They are quick to take flight if approached in the daytime (flying low
and only short distances), but could not really be considered diurnal in habit.
(Preserved = 1-6, 9; photos = lc, 5.)

Miscellaneous Unidentified Sterrhinae

© Genus? sp.?—G.229. N.QLD., Atherton Tableland, 16 mi. ESE of Mareeba, on
a rocky ridge-top 6 mi. up Davies Creek Rd. (D. & NM): A single larva (4 June 72)
on fl. buds of the shrubby Acacia leptostachya Benth. (det. L. Pedley). This most
distinctive small larva could be mistaken for no other; unfortunately it died in the
pupal stage. (Preserved = dry skin of 5, dried and shrunken 6, 7; photos = 5.)

IMMIDAE

e Imma vaticina Meyr. (det. IC)—N.QLD., Clifton Beach, + 14 mi. N of Cairns
(D. & NM): Larvae (9 May 72) on lvs. of the shrub, Fenzlia obtusa Endl.—MYRTA-
CEAE (det. BH). Adults (6, @) emerged 22 May 72; in A.N.I.C., Canberra. See
also Common (1979: 36). (Preserved = 1, 6, 7.)

LASIOCAMPIDAE

e Crexa acedesta Turner (det. IC)—La.20. S.AUST., Blackwood-Belair district
(NM): Captive larvae (March) readily accepted the pendulous mistletoe, Amyema
miquelii, parasitizing Eucalyptus odorata; never on Exocarpos here (det. NM). Adults
fly Feb.—March; rarely seen at uv. light. Sexual dimorphism is fairly notable. See
Common (1966b: 89) for ¢ & @ adult photos. (Preserved = 1-7, 9; photos = 2, 2c,
oH)

48 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

® Crexa punctigera Walk. (det. IC )—La.16 (a synonym may be Sitina anthraxoides
Walk., in B.M.N.H.). S.AUST., Blackwood-Belair district (NM): Larvae (Aug.—
April) fairly common on the shrubby root-parasite, Exocarpos cupressiformis (det.
NM). Adults fly spring—summer; rarely seen at uv. light here. Sexual dimorphism is
notable. See Common (1966b: 89) for larval photo of this sp. at rest on Exocarpos.
(Preserved = 1, 4—7, 8h, 9; photos = 6.)
e Digglesia australasiae (Fab.) (det. IC)—bLa.15. S.AUST., Blackwood-Belair
district (NM): Larvae (most months of the year) on various shrubs, including Acacia
spp. and (especially) Exocarpos cupressiformis (det. NM). Adults (A) fly most
months, with peaks in March—April & July; both sexes frequent at u.v. light (especially
after 2300 hrs). See Common (1966b: 91) or Tillyard (1926: Pl. 30) for ¢
adult photo. (Preserved = 1-6, 9; photos = 1, 2, 5c, 6.)
e Digglesia rufescens (Walk.) (det. SF)—La.18. S.AUST., Blackwood (NM):
Captive larvae (Aug. & Jan.) readily accepted young lvs. of Eucalyptus odorata ( det.
NM). Adults (B) fly Dec.—Feb. & May-July, with peaks in Dec., Feb., & late June—
early July; both sexes frequent at uv. light. See McFarland (1970: 349 & 1972b: 223)
for egg photos. (Preserved = 1-7, 9; photos = 2, 5, 6.)
e Entometa sp. (det. NM)—lLa.14. S.AUST., Blackwood-Belair districts (NM):
Larvae (spring—summer ) on Eucalyptus leucoxylon and E. odorata (dets. NM). Adults
(B) fly March—April (univoltine), esp. after 2300 hrs; both sexes come to uv. light.
Coloration of ¢ & @ nearly identical in this species; forewings are pale tan and hind-
wings are pale orange. A distinguishing feature of this (very large) sp. is its dense
and. tough elliptical cocoon, the smooth outer surface of which is “dyed” bright green
(by a fluid injected into the silk by the larva, during the early phases of cocoon-
construction ); inner cocoon-lining of silk is pure white, however. Regarding the “?”
preceding the reared hymenopterous parasites cited below: These are presumed
from another closely-related Entometa sp., which is the common one in the Adelaide
city area, where these parasite cocoons were collected from a moribund mature larva
at Collinswood, S.AUST., in a city garden (8 Dec. 66, Mrs. R. Smith). See Common
(1966b: 91) for ¢ & ¢ adult photos probably of this sp.; see also Common (1970:
850). (Preserved = 1-7, 8h, 9; photos = 7.)

Opsirhina—see Digglesia.
e Perna exposita (Lewin) (det. IC)—La.19. S.AUST., Blackwood-Belair-Eden Hills
district (NM): Larvae (most months; peak Dec.—Feb.) on mature foliage of Casua-
rina stricta (det. NM). These larvae have only been collected on the tree, C. stricta
in this locality, never on the shrubby C. muelleriana which I have beaten extensively
(all seasons) in search of various larvae. Adults (B) fly spring—autumn; both sexes
frequent at uv. light. See Common (1966b: 91) for ¢ adult photo. (Preserved =
1-7, 9; photos = 2, 5.)
e Perna sp. nov. (det. IC)—La.23. S.AUST., N. Flinders Ranges, 3% mi. S of
Arkaroola Homestead, at roadside (NM): Larvae (30 Oct. 69) on Casuarina sp. (a
tree). The adult is somewhat larger than P. exposita and very much paler, with only
faint (pale tan) maculation on the whitish forewings. (Preserved = 1, 5.)
@ Pinara Pcana Walk. (det. SF )—La.22. S.AUST., Belair, Gloucester Ave. at Ralph
St. (A. Kowanko ): Captive larvae (Sept. 69) readily accepted young lvs. of Eucalyp-
tus odorata (det. NM). Adults fly late winter-autumn; rarely seen at uv. light here.
Sexual dimorphism is notable. See Common (1970: 850) for ¢ adult photo of P. cana.
(Preserved = 1-7; photos = 2, 6.)
e Porela galactodes (Lower) (det. IC)—N.QLD., Atherton Tableland, 12 mi. NE
of Atherton, nr. Tinaroo Pines Caravan Park (+ 2500’) (D. & NM): Larva (late
March 72) on small sapling of the tree, Casuarina littoralis Salisb. (syn. = C. suberosa)
(det. BH). A deformed adult ¢ emerged 16 June 72; in A.N.I.C., Canberra. (Pre-
served = ¢ 1, 5 head capsule only, 6, 7.)

Sitina—see Crexa.

SUPPLEMENT TO VOLUME 33 49

LIMACODIDAE

e Calcarifera ordinata (Butl.) (det. IC)—Lm.7. (1) W.AUST., Moresby Range,
+ 15-20 mi. NNE of Geraldton, in the Oakajee-Howatharra district: Gaudy larvae
(Aug.—mid Oct.) abundant (most years) on 3 spp. of small, low-growing shrubby
acacias here: Acacia ericifolia Benth., A. oxyclada F. Muell. ex Benth., and A. ulicina
Meisn. (dets. B. R. Maslin). These acacias all have small, narrow (linear) phyllodes.
C. ordinata larvae are sometimes so common (locally) as to defoliate their foodplants;
they make no attempt to hide at any time and are most conspicuous. Adults fly mid
Feb.—early April (=); sexes similar in appearance; univoltine. (2) W AUST., Moresby
Range, Howatharra Hill Reserve, + 19 mi. NNE of Geraldton (McFarland, 1977: 19):
The preferred foodplant, in this specific locality, seems to be Acacia ulicina, but A.
ericifolia is also frequently eaten; A. oxyclada is not present on the reserve. (Preserved
= 1, 4-7, 9.) Other records from Howatharra Hill Reserve: 25 Aug. 77 (NM), one
last instar seen feeding on tough, mature stems(!) of Brachysema aphyllum Hook.—
FABACEAE (det. NM); 17 Sept. 77 (NM), 2 last instars feeding on old (tough) lvs.
of Gastrolobium oxylobioides Benth. (Champion Bay Poison Bush! )—FABACEAE
(det. NM), in Zone 3 (SW); 25 Aug. 77 (NM), 2 last instars on mature (tough)
phyllodes of Acacia acuminata in Zone 5 (WC) and 2 feeding on a low, shrubby,
“broom-like” Jacksonia sp. (NM.1175)—FABACEAE, in Zone 4 (C); 28 Sept. 77
(NM), one last instar feeding on leathery, mature phyllodes of Acacia Psaligna Wendl.
(det. NM), in Zone 5 (NE). (3) W.AUST., Northampton, on Lot 351 (Wannere-
nooka Rd.). Larvae (L5—1 Sept. 77) abundant on Acacia tetragonophylla F. Muell.
(det. NM).

e Doratifera oxleyi (Newman) (det. IC)—Lm.5. S.AUST., Blackwood (NM):
Larvae (July—Oct. ) abundant on mature lvs. of Eucalyptus odorata (det. NM); young
larvae scar the If. surfaces with their characteristic “feeding-grooves.” (But see also
the zygaenid, H. tricolor.) Adults (A+) fly March—April only; 9 @ abundant at uv.
light, but ¢ ¢ rarely come to uv. light; univoltine. Peak of ¢ activity probably diurnal
(from + 1000-1500 hrs). Sexual dimorphism is striking in this moth. See Common
(1966b: 67) for ¢ & @ adult photos; Tillyard (1926: Pl. 30) for 6. See McFarland
(1970: 349 & 1972b: 219) for egg photos. (Preserved = 1-3, 5-7, 8dhh, 9; photos
= oe

e Doratifera quadriguttata Walk. (det. SF, IC)—Lm.4. S.AUST., Blackwood
(NM): Larvae (Jan.—Feb.) on mature lvs. of Eucalyptus odorata (det. NM). Adults
(B+) fly Nov.-early Feb. (peak Dec.); sexes similar in appearance and both noc-
turnal; univoltine. (Preserved = 1-5, 9; photos = 2.)

e “Parasa’” sp. (det. IC)—Lm.6. W.AUST., Drummond Cove, +7 mi. N of
Geraldton (NM): Captive larvae (summer) readily accepted mature and semi-mature
Ivs. (phyllodes) of the dominant shrub or low tree of this locality, Acacia ligulata A.
Cunn. ex Benth. (det. B. Maslin). Adults (B+) fly Nov.—-early April (peak Dec.—
Feb.); multiple-brooded; sexes similar in appearance and both nocturnal; @ rarely
taken at uv. light here. Egg very reminiscent of the Pseudanapaea trigona egg. ( Pre-
served = 1-7, 9.)

e Pseudanapaea trigona (Turner) (det. IC)—Lm.3 (a synonym may be P. dentifascia
Hering, in B.M.N.H.). S.AUST., Blackwood (NM): Larvae (early spring—autumn )
on mature lvs. of Eucalyptus odorata (det. NM). Adults (A) fly Oct.—-May (peaks
Noy.—Dec. & Feb.—April); multiple-brooded; sexes similar in appearance and both
nocturnal. The egg is most distinctive with a very soft and entirely transparent chorion.
See McFarland (1970: 349 & 1972b: 219) for egg photos. (Preserved = 1-6, 9;
photos = 1-3, 5-7.)

LYMANTRIIDAE

@ Acyphas leucomelas (Walk.) (det. IC)—Lp.8, 8A, 8B, & 8C. (1) S.AUST.,
Blackwood-Belair district (NM): Larvae (Aug.—Oct.) abundant on young lvs. of

50 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Acacia pycnantha (det. NM). Although they eat only the young lvs., they usually
rest well down the stems on reddish-brown scarred (or partly-damaged and twisted )
mature phyllodes, where their colorful maculation renders them surprisingly incon-
spicuous. Also (less often) found on a variety of other plants, some specific records
being Acacia armata, Pultenaea largiflorens var. latifolia, Exocarpos cupressiformis,
Dodonaea viscosa; also Myoporum viscosum R. Br—MYOPORACEAE and Eucalyptus
odorata (dets. NM). Adults (A) fly Oct-Dec. & Feb.—April; sexes similar in appear-
ance and both come to uv. light. See Common (1966b: 107) for ¢ adult photo of
this or a closely-related sp. (Preserved = 1-6, 8h, 9; photos = @ 1c.) (2) S.
AUST., south coast of Kangaroo Is., 10 mi. W of Vivonne Bay, on land of G. D. Seton
(NM): Larvae (mid Oct. 66) on the low, rounded, “broom-like” root-parasite, Cho-
retrum glomeratum R. Br.—SANTALACEAE (det. MK); also on new lvs. of Platy-
lobium obtusangulum Hook.—FABACEAE (det. NM). (Preserved = 5.) (3) W.
AUST., at Lot 68, Drummond Cove, + 7 mi. N of Geraldton (D. & NM): Larvae
(most months, but especially Aug.—March) abundant and conspicuous on Acacia ligu-
lata A. Cunn. ex Benth. (det. NM); usually on rather thick mature phyllodes, rasping
the surfaces, or partially eating in from the edges. Often on low, dense, windblown
specimens of A. ligulata along the slopes and ridges of the sandhills, close to the beach.
This population may be a distinct subspecies of the eastern leucomelas. (Lp.8C:
preserved = 1, 5-7, 8ddhh, 9.)

Euproctis aliena Butler (Type in B.M.N.H.)—see synonym, A. leucomelas (Walk. ).
e Euproctis marginalis (Walk.) (det. IC)—Lp.11. S.AUST., Blackwood (NM):
Captive larvae (Dec.—Jan.) accepted mature lvs. of Eucalyptus odorata (det. NM).
Nocturnal feeders, probably hiding under loose bark by day. Adults (A) fly Nov.—Jan.
(292 mostly not before late Dec.); sexes similar in appearance and both nocturnal;
2 @ to uv. light mostly before 2300 hrs and ¢ ¢ mostly after; univoltine. See Common
(1966b: 107) for 2 adult photo of a closely-related sp. (Preserved = 1-5, 9; photos
= 19)
e Habrophylla euryzona (Lower) (det. IC)—Lp.12. (1) S.AUST., south coast of
Kangaroo Is., at Seal Bay (NM, M. Pate, & G. D. Seton): Mostly fullgrown larvae
(16 Oct. 66) exceedingly common, crawling over the sand in association with larvae
of my An.6A (Anthelidae) and Ar.34A (Arctiidae); seen eating various low-growing
annual winter herbs (ephemerals), as follows: Crassula sp.—CRASSULACEAE;
Daucus glochidiatus (Labill.) Fisch., Mey., & Avé-Lall., and Hydrocotyle sp.—both
APIACEAE (dets. MK). Adult ¢¢ probably fly from late Oct.—-early Nov.; uni-
voltine. The @ is diurnal, with a peak of activity in mid to late afternoons of sunny,
still days, according to the late J. O. Wilson (pers. comm.). Sexual dimorphism in
this sp. is stupendous! The @ is wingless and highly degenerate. (Preserved = 6 @ 1,
2, 4—4 96, 7, 8dh, 9; photos = $1.) (2) S.AUST., on coast S of Adelaide, Norman-
ville dunes (NM & Mr. & Mrs. J. O. Wilson): Small to half-grown larvae (26 May 69)
abundant, crawling over the dune sand, mostly nr. or under shrubs, where they were
resting and feeding on the small to very small (recently-germinated) seedlings of
winter annuals (various unidentified plant spp. involved ).
® Orgyia anartoides (Walk.) (det. IC)—Lp.9. S.AUST., Adelaide city & suburbs,
and Blackwood (NM): Larvae (most seasons, but especially winter) on many woody
plants. Specific examples: Acacia pycnantha, Hardenbergia violacea, Exocarpos
cupressiformis, *Betula sp. (birch) —-BETULACEAE, and the semi-woody, orange-
flowered *Lantana sp.—VERBENACEAE (dets. NM). The moth is more common
in city & suburbs (garden situations) than in relatively undisturbed areas where the
native flora still predominates. Adult ¢ ¢ (@ degenerate & wingless) fly in spring—
summer (B-— at Blackwood). Sexual dimorphism is spectacular, very similar to the
North American tussock moths (Hemerocampa). (Preserved = 1, 4-7, 9.) See Com-
mon (1966b: 105, 107) for ¢@ & @ adult photos and probably the larva; Tillyard
(1926: Pl. 30) for 3 adult),

SUPPLEMENT TO VOLUME 33 5

NOCTUIDAE

e Achaea janata (L.) (det. NM)—N.125. W.AUST., Exmouth Gulf district
(North West Cape), in the low, coastal sandhills immediately SE of Norcape Lodge:
also in a similar situation + 10-12 mi. S of Exmouth P.O. (D. & NM): Eggs and
larvae of various sizes (7 July 77) on 3 common, low-growing, annual euphorbias,
Chamaesyce sharkéensis (Baillon) Hassall ms., C. australis ssp. glaucescens (Boiss),
and C. coghlanii (F. M. Bail.) Hassall—EUPHORBIACEAE (det. D. Hassall, 1978).
The locality had had one major rain about mid—late May; by early July most annuals
were rapidly maturing (flowering and setting seed). These large, dark larvae made
little attempt to hide and were thus very conspicuous on their foodplants when full-
grown; when small, their dark frass on the fine, white sand was an obvious clue. By
mid July all had pupated; adults emerged Aug. 77. See Common (1966b: 117) for
adult photo. (Preserved = 1, 6.)

e Aedia acronyctoides (Gn.) (det. NM)—N.96. S.AUST., Blackwood-Eden Hills
district (NM): Larvae (Oct. & Jan.) on open, grassy-rocky slopes, on Australian bind-
weed, Convolvulus erubescens Sims—CON VOLVULACEAE (det. MK). Adults (B-)
fly Oct.—April (peaks Nov. & Jan—Feb.). See Common (1966b: 119) for adult photo.
(Preserved = 1, 5, 9.)

© Anomis flava (Fab.) (det. IC)—N.121. N.QLD., Atherton Tableland, Atherton,
at the Forestry Regional Research Station (A. Irvine): Larvae (April 72) abundant
on Hibiscus diversifolius Jacq— MALVACEAE (det. BH). (Preserved = 1, 5, 6, 8d.)
e Buciara bipartita Walk. (det. IC)—N.94. S.AUST., Blackwood-Belair districts
(NM): Larvae (July—early Sept.) nocturmal feeders on new growth of Hibbertia
exutiacies, H. sericea, and H. stricta (dets. MK). Adults (B+) fly late Nov._mid May
(peaks Dec. & April); coming to u.v. light mostly after 2300 hrs, rarely before.

(Preserved — 1 2. 4—7, 9: photos = I, 2.)
e@ Calathusa ischnodes (Tumer) (det. IC)—S.AUST., Aldinga, in + virgin bushland
area (Mr. & Mrs. J. O. Wilson): Larvae (April 67) on Casuarina striata Macklin.
(det. Jf. O. Wilson). Adult (emerged Oct. 67) in the late J. O. Wilson’s collection,
which has recently gone to the A.N.I.C., Canberra. (Preserved = l, 6.)
@ Callopistria maillardi Gn. (det. IC, SF)—N.118. N.QLD., Atherton Tableland,
at Tinaroo Pines Caravan Park (+ 2500’ el.) (D. & NM): Larvae (April-June 72)
in garden on young and semi-mature lvs. of the ornamental fern, *Nephrolepis ?cordi-
folia (var. ?)—POLYPODIACEAE (det. by Limberlost Nursery, Cairns). I am in-
debted to IC for the specific det. and to SF for the generic placement. (Eriopus
Treitschke, 1825, is a junior objective synonym of Callopistria Hiibner, 1821.) It is
of interest to note that the living larvae of 3 Japanese Callopistria spp. are depicted
(with photographs) on various ferns by Mutuura et al. (1970: Pl. 32, figs. 97-99):
of the spp. they figure, C. juventina obscura Butler is fairly close in appearance to
C. maillardi ( especially the adult). (Preserved = 1, 5, 6, 9.)
e Canthylidia zorophanes Turner (det. IC)—N.123. W.AUST., beach areas of
Drummond Cove, +7 mi. N of Geraldton (NM): Larvae (Sept.—Oct.) abundant,
mostly on the staminate ( ¢ ) fls. of the conspicuous beach grass, Spinifex longifolius
R. Br.—POACEAE (det. NM). Adults (A) fly autumn-spring (peak winter). (Pre-
served = I, 5, 6.)

Catephia—see Aedia.

Corrha—see Praxis.
© Cremnophora angasi Walk. (det. NM)—N.116. S.AUST., + 10 mi. NW of Tin-
tinara, at roadside (TN): Larvae (mid Sept. 69) conspicuous on the low-growing
Halgania cyanea Lind|— BORAGINACEAE (det. MK). Reared adult in Newbery
Collection. (Preserved = 5.)
e Dasypodia selenophora Gn. (det. NM)—N.100. S.AUST., Mt. Lofty Range, Stir-
ling West (NM): Captive larvae (Nov.) readily accepted young lvs. of Acacia

52 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

pycnantha (det. NM). Adults recorded for most months except winter. See Common
(1966b: 119) for adult photo. (Preserved = 1-3, 5, 6, 8d, 9.)
e Diatenes igneipicta (Lower) (det. IC)—S.AUST., 5 mi. E of Two Wells (NM):
Larva (19 March 67) beaten from Acacia ligulata A. Cunn. ex Benth. (det. MK).
Adult emerged 12 Dec. 67. (Preserved = 1, 6, 7.)
e Donuca spectabilis Walk. (det. IC)—N.115. W.AUST., Drummond Cove, + 7
mi. N of Geraldton (D. & NM): Captive larvae (Dec. 72) readily accepted lvs.
(phyllodes) of Acacia ligulata A. Cunn. ex Benth. (det. B. Maslin). Adults (B—) fly
spring—summer. (Preserved = 1-6, 9.)
e Earias huegeli Rog. (det. IC)—N.117. W.AUST., 2 mi. S of Kalbarri, Red Bluff
Caravan Park (D. & NM): Larvae (early Nov. 71) on and inside buds, fls., & green
seed capsules of the shrub, Alyogyne hakeifolia (Giord.) Alef—MALVACEAE ( det.
WAH). Adults fly spring-summer. See Common (1970: 863) for adult photo.
(Preserved = 1, 3-7, 8d, 9.)

Eriopus—see Callopistria.
e Eublemma glaucochroa Turn. (det. IC)—N.120. N.QLD., Atherton Tableland,
nr. Tinaroo Pines Caravan Park (+ 2500’ el.) (NM): Larvae (April 72) in “fuzz-
nests” on lvs., and in If. buds and growing tips of the weedy-herbaceous Pterocaulon
sphacelatum Benth. & Hook.—ASTERACEAE (det. BH). (Preserved = 1, 5, 6, 8h,
9.)
e Euplexia dolorosa ( Walk.) (det. IC)—N.119. N.QLD., Atherton Tableland, nr.
Tinaroo Pines Caravan Park (+ 2500’ el.) (NM): Larvae (April 72) on lvs. of
Pterocaulon sphacelatum Benth. & Hook. (det. BH), and P. glandulosum Benth. &
Hook. (det. NM)—ASTERACEAE. The former appears to be preferred in this
locality. (Preserved = 1, 5, 6, 8d, 9.)

Istarva—see Sandava.

Liocola—see Paracrama.

Maurilia—see Paracrama.
@ Meyrickella torquesaria (Lucas) (det. IC)—N.QLD., Atherton Tableland, 8 mi. E
of Mareeba, along rd. to Kuranda (D. & NM): Larva (4 June 72) and pupa (in
cocoon) on native “pine,” Callitris columellaris ssp. intratropica R. T. Bak.—CUPRES-
SACEAE (det. I. Telford). The superb adult ¢ emerged 12 June 72. (Preserved =
1 &; 6,)
e Neumichtis saliaris Gn. (det. SF )—N.104. S.AUST., Blackwood (NM): Captive
larvae (Sept. 66) readily accepted the annual bur clover, *Medicago polymorpha var.
vulgaris (Benth. ) Shinners (det. NM )—FABACEAE. Adults (B) fly spring—summer.
(Preserved = 1-6. )
e Nitocris callimera (Lower) (det. IC)—N.114. (1) W.AUST., south coast at
Eucla ghost town, on white sand dunes (NM & N. B. Tindale): Larvae (Oct.—Nov.
68) always under the sand, usually under or near the bases of small woody shrubs,
where they remain by day. They feed on (unidentified) plant parts, which are just
under the drifting, windblown sand. Several of the common plants of this habitat
probably serve as food. Undoubtedly involved are certain shrubs in the CHENO-
PODIACEAE;; possibly also grasses. The larvae wander about at night, always sub-
surface, leaving concave, meandering trails ( grooves) in the sand, reminiscent of the
common and conspicuous trails made by certain abundant wandering antlion larvae of
this same habitat (Neuroptera: Myrmeleontidae: Acanthaclisus spp.). Adults emerged
after a period of hot days in Jan._Feb. 69; probably univoltine. (Preserved = 1, 4-6,
8dd, 9.) (2) S.AUST., west coast of Eyre Peninsula, nr. Elliston, on white sand dunes
(D. & NM): Larvae (16 Oct. 71) abundant under conditions almost identical to
above; fresh (soft), green frass was under the sand with the resting larvae. Plants
involved not identified.
© Omphaletis norologa (Meyr.) (det. NM)—N.103. S.AUST., + 8 mi. NE of Two
Wells, around borders of sandhills; also nr. Kangaroo Flats, in similar habitat (NM &
TN): Larvae (20, 27 Aug. 66) on the shrub, Rhagodia parabolica R. Br—CHENO-

SUPPLEMENT TO VOLUME 33 53

PODIACEAE (det. MK). Extremely abundant (by beating); preferred are more
dense or compact individuals of the foodplant, especially those in close contact with
the ground. (Preserved = 1, 4-7, 9.)
e Pantydia >capistrata Lucas (det. IC)—W.AUST., Drummond Cove, +7 mi. N of
Geraldton (NM): Larva (May 73) feeding at night on Acacia ligulata (det. B. R.
Maslin). Adult emerged 5 Aug. 73. (Preserved = 1, 6.)
e Pantydia sparsa Gn. (det. IC)—S.AUST., Yorke Peninsula, nr. Cunliffe (NM, N.B.
Tindale, & P. Aitken): Larva (early Nov. 65) on the shrubby root parasite, Exocarpos
aphyllus R. Br. (det. MK). Adult emerged late Nov. 65. See Common (1966b: 119)
for adult photo. ( Preserved = 1, 6, 7.)
e@ Paracrama iocephala (Tumer) (det. IC, SF)—N.108. S.AUST., Adelaide city &
suburbs, especially at Hurtle Square and on the Adelaide Univ. campus (NM): Larvae
(Nov.—June) locally abundant on the N.S.W.-Qld. trees ( planted here as ornamentals ),
*Lagunaria patersonii G. Don—MALVACEAE, and kurrajong, *Brachychiton popul-
neum R. Br—STERCULIACEAE (dets. MK). Adults (B) probably fly most months,
except midwinter; rare at Blackwood. (Preserved = 1, 4-7, 8h, 9; photos = 1, 6.)
e Plusia >argentifera Gn. (det. SF )—N.101 (possibly = subsidens Walk.). S.AUST.,
Adelaide city, at S.A. Museum (NM): Larvae (late May) defoliating ornamental
geraniums, *Pelargonium sp.—GERANIACEAE (det. NM), growing in window-
boxes. (Preserved = 1, 4-7, 8h, 9; photos = 5, 6.)
@ Praxis alterrima (Walk.) (det. IC)—N.102. (This may = synonym of the Lower
ms. name, ~Corrha pandesma.”) S.AUST., Eden Hills, Yalanda St. (orig. 2 from G.
Furness): Captive larvae (Aug.—Sept.) readily accepted new lvs. of Acacia pycnantha
(det. NM). Adults (B-+-) fly late May—Aug. (peak July); univoltine. (Preserved =
17, 9:)

Proteuxoa—see Omphaletis.
© Sandava scitisignata Walk. (det. SF)—N.97. (1) S.AUST., Blackwood (NM):
Captive larvae (Nov. 65) readily accepted and completed growth upon fresh com-
mercial mushrooms, *Psalliota arvensis Schaetf. ex Secr —AGARICACEAE, s.s. (det.
MK). Adults (B—) fly late Sept.early May (peaks Oct., March), coming to u.v. light
esp. after 2300 hrs. (Preserved = 1-5, 9: photos = 1, 2.) (2) S.AUST., south coast
of Kangaroo Is., 10 mi. W of Vivonne Bay (NM): One larva (1 Jan. 66) found under
loose bark of a dying Eucalyptus sp. ( probably eating fungi, or fungal mycelia, which
were present on this tree trunk).

Adults of this sp. have also been taken at u.v. light at Drummond Cove, W.AUST..,
+ 7 mi. N of Geraldton (uncommon).
®@ Spodoptera litura (Fab.) (det. IC)—-W.AUST., Geraldton (NM): Larva (Oct.
73) on tree tobacco, * Nicotiana glauca Grah—SOLANACEAE (det. NM); the record
is of interest because nothing else has been observed feeding on this common weed
here. Adult emerged 25 Nov. 73 at 2140 hrs. (Preserved = 1, 6.)

NOLIDAE

e@ Aquita tactalis (Walk.) (det. IC)—NI1.8 & 8A. (1) W.AUST., 214 mi. S of Kalbarri,
Red Bluff Caravan Park (D. & NM): Captive larvae (Nov. 71) readily accepted
young lvs. of the locally abundant shrub, Melaleuca megacephala F. Muell.—MYRTA-
CEAE (det. WAH). Made typical nolid cocoons of bark surface splinters chewed off
the foodplant stem and woven together from the inside. Adults fly spring—summer—?
(Preserved = 1-7, 9.) (2) S.AUST., south coast of Kangaroo Is., 10 mi. W of
Vivonne Bay, around shore of a small, freshwater lagoon (NM & G. D. Seton): Larva
(mid Oct. 66) on mature lvs. of a tough, dwarfed individual of the shrub, Melaleuca
gibbosa Labill—-MYRTACEAE (det. MK). See Common (1970: 863) for adult
photo. (Preserved = 5, 9.)

Celama—see Nola.

Coesa—see Uraba.

54 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

e Nola bifascialis (Walk.) (det. IC)—N.QLD., Clifton Beach, + 14 mi. N of Cairns
(D. & NM): Larva (9 May 72) on the shrub, Fenzlia obtusa Endl—MYRTACEAE
(det. BH). Adult ( 6) emerged 30 May 72. (Preserved = I, 6, 7.)
e Nola eurrhyncha Turner (det. IC)—NI.4. S.AUST., south coast of Kangaroo Is.,
10 mi. W of Vivonne Bay, around shore of a small, freshwater lagoon (NM & G. D.
Seton): Larvae (13 Oct. 66) on Melaleuca gibbosa Labill. and M. oraria J. M. Black
—MYRTACEAE (dets. MK). The former appears to be the preferred foodplant in
this locality. (Preserved = 1, 5-7, 9.)
e Nola ?lechriopa Hamps. (det. IC)—NI.6 & 6A. S.AUST., Mt. Lofty Range, nr.
Upper Sturt (TN): Larva (24 Sept. 67) beaten from Astroloma conostephioides ( det.
TN). Another record (the same species), beaten from the same foodplant, nr. Norton
Summit (also Mt. Lofty Range), coll. by R. Briggs. (Preserved = 1, 5-7.)
e Nola ?parallacta Meyr. (det. IC)—NI.3 & 3A. S.AUST., Yorke Peninsula, + 2 mi.
S of Kainton (NM, N. B. Tindale, & P. Aitken): Larvae (4 Nov. 65) on the pubescent
undersides of lvs. of the small, open shrub, Olearia pannosa Hook. (det. MK). (Pre-
served = 1, 4-7, 9; photos = lc.) (2) S.AUST., nr. Melrose, on Mt. Remarkable at
2400’ el. (H. M. Cooper): Larvae (24 Sept. 66) on a broad-leafed form of O. pan-
nosa (det. MK). (Preserved = 5.)
e Nola sp. (det. NM)—NI.7. S.AUST., Blackwood (NM): Larva (10 Sept. 67) on
the semi-woody dwarf shrub, Pultenaea largiflorens var. latifolia (det. MK). Preserved
== 5),)
e Nola sp. (det. NM)—W.AUST., Moresby Range, Howatharra Hill Reserve, + 19
mi. NNE of Geraldton (NM): Larva (Nov. 1977) on dormant, immature fl. buds of
Astroloma serratifolium (DC.) Druce (det. WAH). Adult H. 2 Dec. 1977. (Pre-
Senv.edi== IhiGhe)
Roeselia—see Uraba.
e@ Sorocostia hesycha Meyr. (det. IC)—-W.AUST., Drummond Cove sandhills (nr.
beach), + 7 mi. N of Geraldton (NM): Larvae (May 73) on lvs. of Olearia axillaris
(DC.) F. Muell. (det. WAH). Adult emerged 4 June 73. (Preserved = 1, 6.)
@ Uraba lugens (Walk.) (det. SF)—NI.2. S.AUST., Blackwood (NM): Larvae
( July—Oct. & Jan._Feb. ) locally abundant on mature lvs. of peppermint gum, Eucalyp-
tus odorata (det. NM). The smaller larvae are gregarious and skeletonize the lvs. in
a characteristic way. In some years these larvae are so abundant as to nearly defoliate
some of the trees in this locality. (Also on many other Eucalyptus spp.). Adults (A+)
fly Oct.—Dec. & late Feb.—April. This sp. deposits its eggs in groups of almost perfectly
parallel, separated rows. See Tillyard (1926: Pl. 39) for ¢ adult photo; McFarland
(1972b: 229) for egg photos. (Preserved = 1, 2, 4—7, 9; photos = 1, 2, 5, 7.)
e Unidentified large nolid (det. NM)—NI.5. S.AUST., south coast of Kangaroo Is.,
10 mi. W of Vivonne Bay, on the property of G. D. Seton, at the edge of a small
freshwater lagoon (NM & M. Pate): 3 larvae (mid Oct. 66) on tough, dwarfed
individuals (only!) of Melaleuca gibbosa Labill—MYRTACEAE (det. MK). After
return to the mainland, all 3 larvae died; thus no adults were obtained. This record is
included primarily because of the very large size and unique appearance of these
larvae. There are only one or two described nolids that it could be (due to the large
size), and the larvae are most distinctive in morphology and coloration. It was appar-
ently uncommon in this locality, as several hours of beating exclusively M. gibbosa
were required to obtain 3 larvae. (Preserved = 5, 9.)
Zia—see Aquita.
NOTODONTIDAE
(see also Thaumetopoeidae )
@ Antimima corystes Turner (det. IC)—Nd.18. W.AUST., in Kalbarri National Park,
+ 22 mi. E of Kalbarri township, nr. north side of road (NM & N. B. Tindale):
Mature larvae (6 Noy. 68) on young tender tips of the low, rounded, intricate and
spiny perennial, Daviesia hakeoides Meisn., s.l—FABACEAE (det. A. Weston). This
exact locality revisited 19 Nov. 71 (D. & NM); apparently too late, no larvae found

SUPPLEMENT TO VOLUME 33 55

after 30 min of searching, but there were signs of fairly recent feeding on the newer
stem tips and spine-lvs., which were already “hardening off” for the approaching hot
and dry summer. Probably univoltine. (Preserved = 1, 5-7, 9; photos = 1.)

e Antimima cryptica Turner (det. IC, SF)—Nd.19, 19A, & 19B. (1) W.AUST.,
+ 60 mi. SW of Three Springs, on a hill nr. the Hill R. (NM & N. B. Tindale): Larvae
(8 Nov. 68) on soft young lvs. and stem tips of the spiny, prostrate and sprawling,
semi-woody shrub, Jacksonia furcellata (Bonpl.) DC.—FABACEAE (det. WAH).
(Note: Specimens of this same plant det. as J. spinosa (Labill.) R. Br., by plant
taxonomists at Kings Park, Perth). (Preserved = 1, 5, 6.) (2) W.AUST., Darling
Range, + 25 mi. ESE of Lancelin, nr. the Moore R. (NM & N. B. Tindale): Larva
(9 Nov. 68) on the small, spiny, upright dwarf shrub, Jacksonia Psericea Benth. (det.
WAH). (3) W.AUST., Mt. Barren Range, + 6 mi. W of Hopetoun, on top of a
rocky ridge (NM & N. B. Tindale): Larvae (26 Nov. 68) of all sizes (mostly penult.
& last instars) abundant on the upright and leafy shrub, Jacksonia compressa Turcz.
(det. WAH). Probably univoltine. (Preserved = 1, 2, 5, 6, 9; photos = 1, 5.) (4)
W.AUST., 2 mi. S of Kalbarri, nr. Red Bluff Caravan Park (D. & NM): Larva (25
Noy. 71) on the rounded, upright, soft and broom-like, greyish-green shrub, Jacksonia
Plehmannii Meisn. (det. NM).

© Commonia sp. nov. (det. IC)—Nd.21. S.AUST., + 1.5 mi. E of Nundroo (B. &
M. S. Moulds): A single penultimate instar larva (29 Sept. 78) found on Melaleuca
Poraria J. M. Black (det. NM). Mr. Moulds kindly gave me this colorful small larva
during a visit to Drummond Cove, W. Aust.; after its final moult it readily switched to
an offered substitute foodplant, M. uncinata (mature lvs.), upon which it completed
growth without difficulty; fullgrown by 13 Oct. 78 (length 25 mm.); pupated Oct.
17th; adult emerged 2 Nov. 78, at + 2315 hrs; deposited in A.N.I.C. (Preserved =
41, 6,9.)

@ Danima banksiae ( Lew.) (det. NM )—Nd.15, 15A, 15B, 15C, 15D, & 15E. (1) S.
AUST., Mt. Lofty Range, in Belair Nat. Park, 1 mi. E of Belair railway station (NM):
Larvae (24 March 67) on tough mature (sclerophyll) lvs. of the dense woody shrubs,
Hakea rostrata and H. rugosa R. Br. (det. MK). The former is the primary foodplant
in this locality. Adults (B—) fly July—April + (?) (peak Aug.—Sept.). See Common
(1966b: 103) or Tillyard (1926: Pl. 39) for ¢ adult photo; Common (1970: 858) for
line drawing of penultimate instar larva; McFarland (1970: 349 & 1972b: 225) for
egg photos. (Photos = 5c.) (2) S.AUST., Eyre Pen., 10-15 mi. N of Minnipa (C. J.
Winn): Fullgrown larva (16 Sept. 66) on mature (tough-sclerophyll) Ivs. of the
woody shrub, Hakea francisciana FvM. (det. MK). (3) S.AUST., south coast of
Kangaroo Is., 10 mi. W of Vivonne Bay, in dense scrub nr. a freshwater lagoon (NM
& G. D. Seton): Eggs (large and pure chalk-white; very conspicuous) and larvae
(mid Oct. 66) of all instars (mostly smali), abundant on tough, old (sclerophyll) Ivs.
of Hakea muelleriana; also a single larva on old lvs. of the woody, sclerophyll shrub,
Banksia ornata FvM. ex Meisn. (dets. MK). Although Banksia is abundant in this
locality, Hakea is apparently (by far) the preferred foodplant here. Revisiting this
same locality in early Sept. 67, with Dr. E. C. Jaeger, I found fresh adults on the
wing in moderate abundance, coming to uv. light. (Preserved = 1-6, 8h ex egg, 9;
photos = 2.) (4) S.AUST., Mt. Lofty Range, nr. Hahndorf (TN): 5 penult. instar
larvae (6 Dec. 66) on old lvs. of the woody sclerophyll shrub (or small tree), Banksia
marginata (det. NM). (Photos = 5.) (5) S.AUST., inland red-sandy semi-desert,
72 mi. S of Kulgera (N.T.), in a dry creek bed (NM, TN, & L. C. Masterman): 3 eggs
(25 May 67) attached to the long, linear, silvery-gray-pubescent, mature lvs. of a tree
(drooping growth-habit; fls. small & cream-white), probably a Grevillea (det. NM).
See also McFarland (1973: 202). (Preserved = 1, 2, 5, 6; photos = 5.) (6) W.
AUST., 11 mi. SW of Three Springs (NM & N. B. Tindale): Larva (8 Nov. 68) on
tough mature lvs. of the dense and woody sclerophyll shrub, Dryandra cirsioides Meisn.
—PROTEACEAE (det. NM). (Preserved = 5.) (7) W.AUST., Moresby Ranges,
Oakajee-Howatharra district, + 15-20 mi. NNE of Geraldton (NM): Larvae (Aug.—

56 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

early Oct.) frequent on Hakea trifurcata (Sm.) R. Br.; less often on Grevillea pinaster
Meisn. (dets. NM).
e Gallaba stenoptera Turner (det. IC)—“N.” 98. S.AUST., SW Yorke Peninsula,
+5 mi. SW of Carribie Homestead (NM, N. B. Tindale, & P. Aitken): The rather
noctuiform green larvae (2 Nov. 65) were obtained by beating the densely-leafy,
large evergreen shrub, Leucopogon parviflorus (Andr.) Lindl—EKPACRIDACEAE
(det. MK). Probably univoltine. (Incorrect field identification of these larvae orig-
inally caused me to catalog them with the noctuids in my larval collection and notes,
thus the “N.” code-number; the adult determination was not obtained until 6 years
later.) (Preserved = 1, 5, 6, 9; photos = 1.)
e Hylaeora dilucida Felder (det. IC & SF)—Nd.14. S.AUST., Blackwood-Belair
district (NM): Captive larvae (June—Sept.) readily accepted mature lvs. of Eucalyp-
tus odorata; undoubtedly on other Eucalyptus spp. as well. Adults (A) fly March—May
(peak April-early May), coming to uv. light primarily after 2300 hrs; univoltine. See
Common (1970: 857) for ¢ adult photo; McFarland (1970: 350 & 1972b: 225) for
egg photos. (Preserved = 1—7, 9; photos = 2, 5.)
© Neola semiaurata Walk. (det. IC)—Nd.20. AUST. CAP. TERR., base of Black Mt.,
Canberra Botanic Gardens, nr. entrance to exhibit room (D. & NM with R. & S.
Panter): Larvae (16 Jan. 72) defoliating the woody shrub, *Dodonaea multijuga G.
Don.—SAPINDACEAE (det. J. Wrigley). On this date we found 8 last instar larvae,
of large size and in excellent condition, on one small shrub of D. multijuga which is
native to the N.S.W. coast and mts., but not to the Canberra district. See Common
(1966b: 103) for ¢ adult photo and line drawing of last instar. (Preserved = 5, 6, 9.)
Scythrophanes—see Gallaba.
© Sorama bicolor Walk. (det. SF, IC)—Nd.16. S.AUST., Blackwood (NM): Cap-
tive larvae (Jan. 67) readily accepted mature and semi-mature lvs. of Eucalyptus
odorata (det. NM). Adults (B) fly Nov.-Feb. and May—Aug. (peaks Dec.—Jan. &
July), coming to uv. light primarily after 2300 hrs; @ rare at light. Possible evidence
of first instar larval dispersal was noted for this species. See Common (1966b: 105)
for ¢ adult photo. (Preserved = 1-7, 9; photos = 1, 2,5, 6)

OECOPHORIDAE

e Enteremna sp. (det. IC)—-W.AUST., +5 mi. N of Geraldton, along Beatie Rd.
(D. & NM): Larvae (21 Oct. 73) in tough, conspicuous web-frass nests, on mature
(old) lvs. of Banksia attenuata R. Br. and B. menziesii R. Br.-—-PROTEACEAE (dets.
WAH). Adults emerged 23-31 May 74, at night. (Preserved = 1, 6.)

e Myrascia megalocentra Meyr. (det. IC)—Oc.45(M). (1) W.AUST., Moresby
Range, Oakajee district, + 16 mi. NNE of Geraldton (NM & R. G. Swinney): Larvae
(late Aug. 72) abundant in well-formed, individual web-nests, on mature lvs. of Mela-
leuca uncinata R. Br. ex Ait—MYRTACEAE (det. NM); in a dense, undisturbed
heath association, on a rocky hillside. Adults emerged early-mid Nov. 72; deposited
in A.N.I.C., Canberra. (Preserved = 1, 4-7, 8h, 9.) (2) W.AUST., Moresby Range
(+ 600-700’ el.), Howatharra Hill Reserve, + 21 mi. NNE of Geraldton (NM):
Larvae (Aug.—Sept.) often on new lvs. of M. uncinata and M. radula Lindl.; occasion-
ally also on new lvs. of M. megacephala F. Muell and M. scabra R. Br. (s.l.); all of
these are relatively dense woody shrubs. See Common (1977); also Common & Bellas
(1977).

© Thudaca obliquella Walk. (det. D. J. Carter) —Oc.44(M). S.AUST., Mt. Lofty
Range, Belair Nat. Park, 1 mi. E of Belair railway station (NM & D. Bakker); Larvae
(22 Sept. 69) abundant on tough mature (sclerophyll) Ivs. of Leptospermum myr-
sinoides (det. NM). See Tillyard (1926: Pl. 28) for adult photos. (Preserved = 1,
!-6; photos = 6.)

PTEROPHORIDAE

© Trichoptilus sp. (det. I1C)—W.AUST., Drummond Cove sandhills nr. beach, + 7
mi. N of Geraldton (NM): Larvae (Nov.—Dec. 74) on (only) fl. buds, fls., and

\
J

SUPPLEMENT TO VOLUME 33 57

green frs. of Boerhavia chinensis (L.) Aschers. & Schw.—NYCTAGINACEAE ( det.
WAH). The pupae took only 5-6 days to hatch! Adults emerged 7-15 Dec. 74.
(Preserved = 1, 6.)

PYRALIDAE
(A) SUBFAMILY EPIPASCHTIINAE

e Epipaschia (Macalla) pyrastis (Meyr.) (det. IC, D. J. Carter)—Py.31(M). _ S.
AUST., Blackwood (NM): Larva (Feb. 67) in tubular and extended web-nest, among
lvs. of a small sapling of Eucalyptus leucoxylon (det. NM). A series of larvae, reared
by the late J. O. Wilson (from eggs of a Blackwood @, April 68), readily accepted
mature lvs. of E. odorata and reached last instar in early July. Adults (B—) fly spring—
autumn. See McFarland (1970: 349 & 1972b: 223) for egg photos. (Preserved = 1,
2. 2. 6, 9: photos — 1.)

(B) SupFAMILY PyRAUSTINAE

e *Mecyna polygonalis Hbn. (?) (det. NM)—Py.23(M)A. (1) S.AUST., Adelaide
suburbs and Blackwood (NM): Larvae (Nov.) often on *Genista maderensis; also
reported (by M. Boyce), in a garden at Burnside, on young lvs. of a South Australian
native sclerophyll, Hovea longifolia var. lanceolata (Sims) Benth., and on *Podalyria
sp.—all FABACEAE (dets. NM). Adults (B+) fly spring—autumn. (Preserved = 5,
6.) (2) W.AUST., +7 mi. N of Geraldton, at Drummond Cove (D. & NM): Larvae
(Sept.—Oct.) often defoliate the coastal shrub, Templetonia retusa (Vent.) R. Br. ex
Ait.— FABACEAE (det. NM). (3) W.AUST., Moresby Range, Howatharra Hill
Reserve, + 19 mi. N of Geraldton (D. & NM): Larvae (June-July) feeding on lvs.
of the small native shrub, Bossiaea biloba Benth— FABACEAE (det. NM).

SATURNIIDAE

e Antheraea helena (White) (det. NM)—St.21. (1) S.AUST., Blackwood (NM):
Captive larvae (Nov.—Dec. 66) readily accepted young lvs. of Eucalyptus odorata
(det. NM). Adults (B) fly late Sept.—March (peaks Oct.-Nov. & Feb.); univoltine.
See Common (1970: 850) for ¢ adult photo. (Preserved = 1-7, 9; photos = 3c, 4c,
5c.) (2) S.AUST., Naracoorte, nr. Lochiel Ave. (NM & TN): A newly-moulted last
instar larva (27 Dec. 68) on luxuriant semi-mature (sclerophyll) lvs. of a sapling of
Eucalyptus baxteri (Benth.) Maiden & Blakely ex Black (det. MK). (3) S.AUST.,
Mt. Lofty Range, Stirling, in a nursery (NM): A full-grown and healthy last instar
larva (2 Feb. 67) defoliating a small, ornamental weeping birch tree, *Betula sp.—
BETULACEAE (det. NM).

SPHINGIDAE

@ Agrius convoluuli (L.) (det. NM)—Sp.22. S.AUST., Campbelltown (Adelaide
suburb): Last instar larvae (early—mid May 66) in several gardens, on the ornamental,
blue-fl., annual morning glory, *Convolvulus sp—CONVOLVULACEAE (det. NM).
Green, brownish, and black phase larvae were present. Adults (C) fly spring—summer
at Blackwood; to light especially after 2300 hrs. See Common (1966b: 99) for Q
adult photo. (Preserved = 5, 6, 9.)

Celerio—see Hyles.

Herse—see Agrius.
@ Hippotion celerio (L.) (det. NM)—Sp.21. (1) S.AUST., Northfield (Adelaide
suburb), at 19 Wright Ave. (D. Daulby): Larvae (28 March 66) defoliating the
(African) arum “lily,” *Zantedeschia aethiopica (1...) Spreng —ARACEAE (det. NM).
These larvae were full size (last instar) and thriving on that plant. (Preserved = 1, 5,
6, 9.) (2) S.AUST., Blackwood (NM): Larvae (spring—autumn) on grape vines,

58 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

*Vitis sp —VITACEAE, which is the usual foodplant here. Adults (B-—) fly late
spring to mid May. (3) W.AUST., Moresby Range, Howatharra Hill Reserve, + 19
mi. NNE of Geraldton (D. & NM & E. C. Jaeger): Larvae (July—Sept.) on
Clematicissus angustissima (F. Muell.) Planch VITACEAE (det. NM), a vine which
spreads over the ground and shrubs. (4) W.AUST., +7 mi. N of Geraldton, Drum-
mond Cove (D. & NM): Larvae (Oct.—-Nov.) on Boerhavia chinensis (L.) Aschers.
& Schw.—NYCTAGINACEAE (det. P. Wilson), a perennial sp. growing on coastal
sandhills. (5) W.AUST., 9 mi. S of Carnarvon, at roadside (D. & NM): Larvae (12
July 77) in abundance on the noxious weed, *Emex australis Steinh.—POLYGON-
ACEAE (det. NM); they were eating this in preference to many other weeds and
native annuals growing here profusely, in response to rains about 6—7 weeks earlier.
e Hippotion scrofa (L.) (det. NM)—Sp.20. S.AUST., Goodwood (Adelaide sub-
urb): Larvae (March 66) on *Fuchsia sp.—ONAGRACEAE, and *Coprosma baueri
Endl.— RUBIACEAE (dets. MK). Adults (A) fly late Aug.—April (peaks Nov. &
Feb. ), coming to uv. light mostly after 2300 hrs (Blackwood, S. AUST). See Common
(1970: 857) for @ adult photo. (Preserved = 1, 5, 6, 9; photos = 5, 5c, 6c.)

® Hyles lineata livornicoides (Lucas) (det. NM )—Sp.23 & 23B. (1) N.TERR., 56
mi. S of Alice Springs (NM): Larvae (5 April 66) very abundant, on luxuriant spread-
ing “mats” of an annual Boerhavia sp—NYCTAGINACEAE (det. NM); most were
penult. or last instar on this date. (It was obvious that this locality probably had
received heavy rains during hot weather in Jan., Feb., or early March, which would
account for the luxuriance of this plant and for the presence of many large larvae in
early April.) See Common (1966b: 101) for 9 adult photo. (Preserved = 5, 9.)
(2) S.AUST., Blackwood (NM): Captive larvae (March 68) at first accepted the
ornamental garden annual, *Mirabilis jalapa L.—NYCTAGINACEAE (det. MK), but
did not thrive. Adults (C) fly Feb.—_March (coming to uv. light mostly after 2300 hrs)
and appear only sporadically in the Blackwood district; they were only seen one year
(1968) out of the 5 summers (1965-1969) that I lived there, but several came to light
that summer, and they were also reported from other Adelaide suburbs in early 1968.
(Preserved = 1-4, 9.) (3) S.AUST., Hallett Cove (S of Adelaide), along the cliff
edge to the south of the cove beach (NM): Mature larvae (late March 68) were
present on scattered individuals of Boerhavia diffusa L. (det. NM). (There had been
unusual heavy summer rains in the Adelaide vicinity during hot weather in Jan.—Feb.

68 ).

THAUMETOPOEIDAE

@ Cynosarga ornata Walk. (det. IC)—Ta.13. N.QLD., Atherton Tableland, around
gravel quarry nr. Tinaroo Pines Caravan Park (+ 2500’ el.) (D. & NM): Larvae
(March—June ) of all sizes and unhatched eggs, very conspicuous on small saplings of
the tree, Casuarina littoralis Salisb. (synonym = C. suberosa) (det. BH). These larvae
are highly gregarious in early instars, retaining this behavior (to a very slight degree )
even into last instar. Adults (March—June) were never attracted to uv., mercury vapor,
or ordinary incandescent light (including the hours from midnight to dawn) during
this period, nor did we ever see them flying in the daytime. However, they were
regularly emerging (in captivity) all through April-June. (Preserved = 1-7, 8h, 9.)
@ Discophlebia catocalina Felder & Rogenhf. (det. SF, IC)—Ta.3. S.AUST., Black-
wood (NM): Captive larvae (Jan._Feb.) readily accepted mature (tough) lvs. of
Eucalyptus odorata (det. NM); strictly nocturnal feeders when past the early instars.
First instar larval dispersal is notable in this species. After dispersal, they still retain
semi-gregarious tendencies when small, but later become solitary. Adults (B+) fly
mid Noy.—March (peak late Dec.—early Feb.), coming to lights especially on hot
nights; univoltine. See McFarland (1970: 350 & 1972b: 227) for egg photos. Pre-
served = 1-7, 9; photos = lI, 2.)

© /picoma argentata (Walk.) (det. IC)—Ta.6. N.TERR., 27 mi. E of Timber Creek
store (NM): 16 half-grown gregarious larvae (18 April 66) at rest, closely side-by-

SUPPLEMENT TO VOLUME 33 59

side, all on one mature If. (of a small sapling) of an unidentified Eucalyptus sp. (det.
NM), which was the predominant eucalypt in that locality. These larvae were trans-
ported 2,000 mi. to the south, readily accepting mature lvs. of various eucalypts
enroute; they were then reared to pupation on mature lvs. of E. odorata (which they
readily accepted) at Blackwood. (They refused to feed unless kept warm, however,
as it was early winter in the south, and they had been transported from the tropics. )
(Preserved = 1, 4—7, 8h, 9; photos = 5.)

e Epicoma melanosticta Donovan (det. IC, SF )—Ta.5; Ta.11. (1) S.AUST., Black-
wood (NM): Captive larvae (April—July 66) readily accepted old (mature) sclero-
phyll lvs. of Eucalyptus odorata (det. NM) in captivity, feeding only or mostly at
night when older; highly gregarious in early instars, but gradually becoming essentially
solitary by last instar. I have 3 separate records of field-collected larvae of this sp.
feeding on E. odorata, on 22 Jan. 67, 5 Feb. 67, and 27 Sept. 68; in two of the cases
the larvae were small, resting or feeding in intimate aggregations of between 20 and
30 individuals; they were on very small saplings of the foodplant, not far above the
ground. Adults (B) fly Nov.—mid June (peaks Dec. & late March—-early May); flight
nocturnal (occasionally crepuscular), both sexes coming readily to uv. light. See
Common (1970: 857) for ¢ adult photo; McFarland (1970: 350 & 1972b: 225) for
egg photos. (Preserved = 1-7, 9; photos = 1, Ic, 2, 5.) (2) W.AUST., Mt. Barren
Range, + 6 mi. W of Hopetoun, on top of a rocky ridge (NM & N. B. Tindale): Two
last instar larvae (26 Nov. 68) feeding (0600-0700 hrs), close to the ground, on old
lvs. of the shrub, Calothamnus validus S$. Moore—MYRTACEAE (det. WAH). AI-
though I recorded these larvae under a separate code-number (Ta.1l1), a 2 adult
(H. 11 Jan. 69) later proved to be Epicoma melanosticta. (Preserved = 1, 5, 6, 9;
photos = 1, 5, 6.) (3) W.AUST., Moresby Range, Howatharra Hill Reserve, + 19
mi. NNE of Geraldton (NM & Lisa Green): 3 aggregations of half-grown larvae (12
Aug. 78) on old lvs. of Calothamnus homalophyllus F. Muell—MYRTACEAE ( det.
NM); in Zone 4(C) & (WC) of the reserve. (This is probably also “Ta.11.’’)

e Epicoma (sp. nov.?), nr. tristis Lewin (det. NM) or nr. melanospila (Wallengren )
(det. IC)—Ta.4. S.AUST., south coast of Kangaroo Is., 10 mi. W of Vivonne Bay,
on land owned by G. D. Seton, around the edges of a small freshwater lagoon (NM &
M. Pate): Larvae (31 Dec. 65 and 30 Jan. 66) uncommon and scattered, on various
sclerophyll dwarf shrubs of the habitat (a rather dense and richly-varied heath asso-
ciation), but not on eucalypts. Specifically seen feeding on Darwinia micropetala
(FvyM.) Benth MYRTACEAE (det. MK). Larval feeding and activity was diurnal.
Adults probably fly April-May; a single most distinctive ¢ adult emerged (in a heated
room indoors) on 7 May 66. I have not seen the @. The adults of this sp. may be
both diurnal and nocturnal in activity; although I suspect the former, available evi-
dence points in both directions! In my opinion this sp. shows a closer kinship with
Epicoma tristis (both in the larval stage & adult) than with E. melanospila, although
it is easily separated from either of these. See Common (1966b: 105) for ¢ & Q adult
photos of E. melanospila. (Preserved = 61,5, 6; photos = ¢ 1.)

e Epicoma tristis Lewin (det. IC, SF)—Ta.8 & 8A. (1) S.AUST., Belair Nat. Park,
1 mi. E of Belair railway station, just S of Sheoak Rd. (NM): Larvae (Oct.—Feb. )
regularly on two completely unrelated shrubs, Casuarina muelleriana and Leptosper-
mum myrsinoides (dets. NM). The former appears to be the “preferred” foodplant in
this locality, although both are readily eaten in captivity. The larvae are gregarious
when small, but become solitary and widely-scattered in later instars. They feed
diurnally; sunlight and/or warmth are needed to stimulate optimum feeding activity.
Adults (A), of very patchy and localized distribution, fly March-April (peak short;
usually late March—early April only); strictly diurnal in activity, with ¢ flight entirely
on mornings of warm days, between + 0800-1130 hrs (peak 0815-1000 hrs) when
they seek the slightly smaller 2 2 resting in low bushes of the “colony” or population-
center; univoltine. (Preserved = 1-7, 9-extensive; photos = 1, 2, 5, 6.) (2) S.
AUST., 3.3 mi. SE of Blackwood Post Office (NM): Larvae (spring—-summer) were

60 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

entirely on the abundant Leptospermum myrsinoides (det. NM) in this population.
(For remarks on diapause in the egg stage of this sp., see McFarland, 1973: 199.)
Ochrogaster—see Teara.
@ Oenosandra boisduvalii Newman (det. SF, IC)—Ta.1. (The “r’ belongs in the
generic name: letter from SF, 25 Aug. 1971.) S. AUST., Blackwood (NM): Captive
larvae (June-Sept.) readily accepted mature lvs. of Eucalyptus odorata (det. NM);
strictly nocturnal feeders, hiding by day under loose bark on the eucalypt trunks and
branches. First instar larval dispersal is notable in this sp., but after dispersal they
still show semi-gregarious behavior (resting closely side-by-side under the bark).
Capable of rapid locomotion to and from feeding and resting sites, which are
often far apart. Their behavior, in this respect, is much like that of the North
American arctiid, Hemihyalea edwardsi Pack. Adults (A) fly mid March—-mid May,
with ¢ ¢ coming to uv. light mostly after 2300 hrs; univoltine. Sexual dimorphism
in this sp. (both color & maculation) is very striking; the sexes are often widely
separated in collections! See McFarland (1970: 350 & 1972b: 227) for egg photos.
(Preserved = 1-6, 9; photos = 1, Ic, 2.)
e Teara ?contraria (Walk.) (det. NM )—Ta.7 & 7A-7D. (1) S.AUST., along road
to Woomera, NW of Port Augusta (NM): Larval nest-bags (3 April 66) present in
countless numbers on mulga trees, Acacia aneura FvM. ex Benth. (det. NM); some
single trees contained several dozen large (last instar) larval nests. Many of these nests
were occupied by nearly fullgrown larvae (Ta.7). For numerous details on the early
stages of this species and the following (Ta.9), which may be a separate species, see
the interesting W.Aust. observations by Mills (1951-52). (2) S.AUST., inland
desert, 64 mi. S of Coober Pedy (NM): Larvae (4 April 66) abundant, inside their
large and conspicuous, + spherical or elliptical whitish silken bag-nests, on Cassia
nemophila var. platypoda R. Br. (Benth.) CAESALPINIACEAE (det. MK). The
larvae are primarily nocturnal feeders, and highly gregarious at all times for the entire
duration of the larval stage. They rest by day inside the nest-bags, which are rather
fully packed with dried larval frass; the frass no doubt plays an important part in
temperature-regulation within the nest. Just prior to pupation (autumn to early win-
ter) they leave the nests and wander long distances over the ground, in characteristic
single-file processions, searching for pupation-sites; these “larval ropes” are often seen
crossing roads in the autumn (April +). Adults fly spring—early summer; crepuscular
and nocturnal, both sexes (but mostly ¢ ¢ ) coming abundantly to uv. light; the 2 @
sometimes fly in late afternoon, but not in great numbers; univoltine. (Preserved = 1,
2, 5, 6, 8d, 9; photos = 5c, 7c-nests.) (3) S.AUST., 44 mi. S of Coober Pedy, in a
dry creek bed (NM): Larvae (4 April 66), of what I suspect are probably this same
species, in silk nests on the branches, and in trunk crotches of an unidentified Eucalyp-
tus sp. (det. NM). Some of the trees in this locality were nearly defoliated at the
time (Ta.7B).
e Teara sp. (det. NV)—Ta9. (1) S.AUST., Eyre Peninsula, + 5 mi. N of Streaky
Bay (R. Edwards): Half-grown larvae (mid Feb. 67) on the coastal Acacia anceps
DC. (det. MK); upon capture they were crawling down the trunk in a single-file
procession (sunny morning at 0730 hrs). These larvae were reared to full size (late
April) at Blackwood, S. Aust. on a readily-accepted substitute, Acacia pycnantha.
They did not make any nest-bags, but rested in a heap, all piled up together, on or in
loose litter at or near the base of the foodplant, usually among their cast larval skins,
on somewhat of a slight silken mat covering the litter. They were sporadic nocturnal
feeders, on some nights not feeding at all. It would be interesting to know exactly
what factors determine the nights of feeding. (Preserved = 1, 4-6, 9.) (2) S.AUST.,
at base of Black Hill, Athelstone, E of Adelaide (TN): About 30 small larvae, of +
20-25 mm length (15 Jan. 67), sitting heaped together in a “clump,” covering part
of a If. and stem, up on a branch of Acacia pycnantha Benth MIMOSACEAE, ( det.
TN). There was no sign of a bag (or any other silken nest-like structure) anywhere
on this plant or nearby. I have not seen this particular series of larvae, but suspect
they are identical to my Ta.9, based on the information provided by Mr. Newbery.

SUPPLEMENT TO VOLUME 33 61

Regarding the early stages of what is probably Ta.9 in Western Australia, see Mills
(1950-52). There exists some question as to whether or not this sp. (Ta.9) and the
foregoing (Ta.7) are, in fact, two distinct spp.; I am inclined to agree with Mills
(1951: 61, 66-67 and 1952: 87-92) that they are, but this opinion is based only upon
observations of the larval stages. Aside from striking behavioral differences, the longest,
silk-soft, pale gray hairs on my Ta.9 larvae were notably longer than were the longest
hairs of the bag-forming Ta.7 larvae. (If both Ta.7 & Ta.9 are in fact one and the
same species, as stated by some authorities, then it is an incredibly plastic species,
having not only two morphologically distinct larval forms, but also two differing
patterns of behavior and habits. )

Trichetra—see Trichiocercus.

e Trichiocercus sparshalli (Curtis) (det. IC, SF)—Ta.2. S.AUST., Blackwood
(NM): Captive larvae (Jan._Feb.) readily accepted mature lvs. of Eucalyptus odorata
(det. NM). These larvae are highly gregarious when small. This behavior gradually
weakens as they grow larger, but is still retained (to some degree) into last instar.
The morphology of certain structures associated with the larval mouth parts is quite
peculiar and warrants close investigation. Adults (A) fly late Oct.-mid May (peaks
Noy.—Jan. & March), coming to uv. light especially after 2300 hrs. Both sexes are
strictly nocturnal, but the @ is rarely seen at lights here. The ¢ is entirely pure chalk-
white, including all parts of the abdomen; in the @, the dense, terminal, abdominal
tuft of deciduous scales is light golden-tan or golden-brown. (Preserved = 1-7, 9;
pnotos — 1, 2.5, 5c.)

e Trichiocercus sp. nov. (det. IC, SF)—Ta.10. S.AUST., Blackwood (NM): Cap-
tive larvae (July—Sept. 68) readily accepted mature Ivs. of Eucalyptus odorata ( det.
NM). Larval and adult behavior is much as described for Trichiocercus sparshalli,
but larval appearance and hair coloration in these two spp. is very strikingly different.
Adults (B—) fly only from late April—early June, coming to uv. light mostly after 2300
hrs; univoltine. They are, on the average, somewhat larger than T. sparshalli adults,
and both sexes are marked by a variable zone (not sharply defined) of sooty-blackish
coloration on the abdominal dorsum. In the @, the dense terminal tuft is primarily
black with an overlay of longer, hair-like, grayish-brown scales. (Preserved = 1-7, 9;
photos — 1 2 4, Ac. 5, 6.)

XYLORYCTIDAE

e Cryptophasa melanostigma (Wallengren) (det. IC)—N.QLD., Atherton Tableland,
around gravel quarry nr. Tinaroo Pines Caravan Park (+ 2500’ el.) (N. B. Tindale,
collector): Larvae (May-June 72) in short tunnels inside stems of Casuarina littoralis
Salisb. (synonym = C. suberosa) (det. BH). Adults emerged about Sept.—Oct. (Pre-
served = 1, 6.)

® Cryptophasa sp. (det. NM)—W.AUST., Drummond Cove, + 7 mi. N of Geraldton
(D. & NM): Larva (Aug.—Sept. 76) in short tunnel inside a lower branch-stem of a
young (2-year-old) Acacia ligulata A. Cunn. ex Benth. (det. NM). An adult (with
cream-tan forewings ) emerged in Oct. 76.

e Lichenaula sp. (det. IC)—Xy.41(M). W.AUST., +4 mi. N of Kununoppin, on
top of a huge granite dome, Waddouring Hill (NM & N. B. Tindale): Larvae (1 Nov.
68 ) inside frass-covered silken tubes among lichens, on the granite surface, and feeding
(at night and perhaps on cloudy days) on the (unidentified) LICHENS surrounding
them. Several adults emerged, from the field-collected larvae, in March—April 69.
(Preserved = 1, 5, 6.)

© Genus? sp.? (det. IC)—47(M). W.AUST., Drummond Cove, +7 mi. N of Gerald-
ton (D. & NM): Larvae (March 74) in web-tunnels beneath the prickly dwarf shrub,
Acanthocarpus preissii Lehm.—XANTHORRHOEACEAE (det. WAH); on the sand-
hills immediately behind the beach. This record is included primarily for the unusual
foodplant involved. The larval nests are conspicuous tangled webs inside the lower

62 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

parts of the plant, with a tough silken tube extending several inches down into the
fine sand beneath the plant. The larvae remain in the tube (below ground) by day,
coming out to feed on the tough lIvs. after dark. Adults emerged June-July 74. The
2 has oddly-narrowed, reduced wings, looking deformed. (Preserved = 1, 5, 6.)

ZYGAENIDAE

e Hestiochora rufiventris (Walk.) (det. SF, IC)—Zy.8 & 8A. (1) S.AUST., +5
mi. S of Monarto South, at Chauncey’s Line (M. Fagg): Larvae (2 Oct. 66) abundant
on tough mature lvs. of a single bush of Melaleuca sp—MYRTACEAE (det. M.
Fagg). This is presumed to be M. lanceolata Otto, which grew in that locality (det.
NM). (Preserved = 1, 4, 5, 6, 9; photos = 1.) (2) W.AUST., south coast, on
rim of escarpment + 43 mi. W of Eucla (NM & N. B. Tindale): Adults and fresh
eggs (2 Dec. 68) extremely abundant on large shrubs of Melaleuca lanceolata Otto
(det. WAH). The adults are weak fliers and diurnal; probably univoltine. See Mc-
Farland (1970: 349 & 1972b: 219) for egg photos. (Preserved = 1, 2, 3, 9; photos
— im)

e Hestiochora tricolor (Walk.) (det. SF, IC)—Zy.9. (1) S.AUST., +3 mi. S of
Port Willunga, at the north edge of the “Aldinga Scrub,” just S of Fraser St. at Bristol
St. (D. & NM with the late Mr. & Mrs. J. O. Wilson): Nearly fullgrown but dormant
larvae (28 July 70) resting on mature lvs. of a mallee eucalypt, Eucalyptus fasciculosa
(det. MK), and on one other unidentified Eucalyptus sp. The characteristic larval
“feeding-grooves,” on the mature leaf surfaces, are easily recognized. (However, see
also the limacodid, Doratifera.) Earlier in the year (14 March 70; D. & NM), freshly-
emerged diurnal adults were on the wing in this same locality. Indications are that
populations of this moth (a weak flier) tend to be restricted to very localized “col-
onies’; this remark may also apply to H. rufiventris. See Common (1966b: 65) or
Tillyard (1926: Pl. 28) for adult photos. (Preserved = 1-7, 9; photos = 1-3.) (2)
W.AUST., Kalbarri Nat. Park, + 23 mi. E of Kalbarri township, nr. picnic area at
Hawks Head Lookout, overlooking south side of Murchison River (D. & NM with K.
& L. Haines): Adult 2 sighted (mid-day, 23 Sept. 74), but I was not able to capture
it. Inspection of the leathery, mature lvs. on the small mallee eucalypts here revealed
old, dry scars caused by groove-like “feeding-tracks,” probably made earlier in the
year by larvae of this species. (This may be a W. Aust. ssp. of H. tricolor.)

e Pollanisus apicalis Walk. (det. SF, IC)—Zy.6. S.AUST., +8 mi. NE of Two
Wells, on white sandhills (NM, TN, & M. Pate): Larvae (20, 27 Aug. 66) locally
abundant on buds, fls., & young lvs. of scattered individuals of the dwarf shrub,
Hibbertia virgata R. Br. ex DC. (det. MK). Adults (Oct.) diurnal, weak fliers; prob-
ably univoltine. (Preserved = 1, 4—6; photos = 1, 5c.)

e Pollanisus dolens Walk. (det. SF)—Zy.7. S.AUST., Mt. Lofty Range, nr. Long-
wood, at Aldgate (Heather Rd.), and in Belair Nat. Park, 1 mi. E of Belair Railway
Station (NM & TN): Larvae (Aug.—Sept.) common on tough, mature lvs. of Lepto-
spermum myrsinoides (det. NM). Adults (Oct.) diurnal; weak fliers; probably uni-
voltine. (Preserved = 1, 4—7, 8d, 9; photos = 1, 5.)

e Pollanisus viridipulverulentus Guérin (det. NM, SF)—Zy.5. S.AUST., Blackwood-
Belair district (NM): Larvae (July—Aug.) on buds, fls., and young lvs. of Hibbertia
stricta and H. sericea (dets. MK); the former appears to be preferred around Black-
wood. Adults (B+) fly late Sept.—Oct., occurring in localized colonies; weak fliers
and diurnal; probably univoltine. See Common (1966b: 65) for ¢ & @ adult photos.
(Preserved = 1, 5-7, 8d, 9; photos = 1.)

SUPPLEMENT: Fijr ISLANDS

AGARISTIDAE

@ (?)Sarbanissa (Seudyra) bostrychonota Tams (det. H. S. Robinson)—As.10. VITI
LEVU, nr. Nadarivatu, along trail about halfway up to the lookout on Lomalagi Peak

SUPPLEMENT TO VOLUME 33 63

(NM & G. F. Gross): Two larvae (15 Feb. 68) feeding on the non-urticating lvs. of
Leucosyke corymbulosa (Wedd.) Wedd.—URTICACEAE (det. J. W. Parham). Adults
are tentatively det. as above, even though none were reared out, because Robinson
(letter of 23 July 68) stated that “. . . this is the only agaristid recorded as occurring
here...” See Robinson (1975: 111 & Pl. Figs. 10-11) for ¢ and @ adult photos.
(Preserved = 5, 9.)

ARCTIIDAE
SUBFAMILY NYCTEMERINAE

e Nyctemera baulus (Bdv.) (det. NM)—Ar.40. VITI LEVU, Navai-Nadarivatu
district, in weedy cornfields (NM): Adults, fresh eggs, and larvae of all sizes abundant
(10 Feb. 68), on the rank and weedy annual, *Crassocephalum crepidioides ( Benth. )
J. Moore—ASTERACEAE (det. J. W. Parham); I also saw 2 larvae, on this same
foodplant in a private garden in Suva (late Feb. 68). See Robinson (1975: 107 &
Pl. Fig. 9) for ¢ adult photo. For egg photos of a close relative, N. amica (White),
see McFarland (1970: 350 & 1972b: 229).

REFERENCES
BOTANICAL:
Bearp, J. S. 1965. Wildflowers of the Northwest [Australia]. Westviews, Perth.
28 pp.

1969. Endemism in the Western Australian flora at the species level. J.

Roy. Soc. W. Aust. 52(1): 18-20.

(ed.). 1970. A descriptive catalogue of West Australian plants (2nd ed.).

Society for Growing Australian Plants, Sydney.

& A. C. Burns. 1976. Vegetation survey of Western Australia—The vegeta-
tion of the Geraldton area [map and text]. Vegmap Publications, 6 Fraser Rd.,
Applecross, W. Aust. 6153.

BLACKALL, W. E. & B. J. Grieve. 1974. How to know Western Australian wild-
flowers, Parts I-III [1974 ed. reprinted in one vol.]. Univ. of W. Aust. Press,
Perth. [Part IV, see Grieve. ]

Biack, J. M. 1943-1957. Flora of South Australia, Parts I-IV (2nd ed.). Govt.
Printer, Adelaide. [See also Eichler. |

Bursipce, N. T. 1963. Dictionary of Australian plant genera (Gymnosperms and
Angiosperms ). Angus & Robertson, Sydney.

CLELAND, J. B. 1932-34. The flora between Outer Harbour and Sellick’s Beach,
South Australia [Adelaide district]. S. Aust. Nat. 14: 45-48, 55-56, 109-120; 15:
9-13, 33-40.

Corron, B. C. (ed.). 1964. South Australian National Parks and Reserves. Govt.
Printer, Adelaide. [Belair National Park covered in detail. ]

ErcHLer, Hy. 1965. Supplement to J. M. Black’s Flora of S. Aust. Govt. Printer,
Adelaide.

Erickson, R., A. S. Georce, N. G. MarcHant, & M. K. Morcomset. 1973. Flowers
and plants of Western Australia. A. H. & A. W. Reed, Sydney.

Francis, W. D. 1970. Australian rain-forest trees (3rd ed.). Aust. Govt. Publishing
Service, Canberra.

Garpner, C. A. 1944. The vegetation of Western Australia with special reference
to the climate and soils. J. Roy. Soc. W. Aust. 28: xi-Ixxxvii [= Presidential
Address, 1942].

1959. Wildflowers of Western Australia. W. Aust. Newspapers, Ltd.,
Perth.

Grieve, B. J. & W. E. Buackaty. 1975. How to know Western Australian wild-
flowers, Part IV. Univ. of W. A. Press, Perth.

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Haut, N., R. D. JoHNsron, & G. M. CHIPPENDALE. 1970. Forest trees of Australia
(3rd ed.). Aust. Govt. Publishing Service, Canberra.

Lamp, C. & F. Cotter. 1976. A field guide to weeds in Australia. Inkata Press,
Melbourne.

McFartanp, N. 1970. Botanical names in entomological papers and habitat studies.
J. Res. Lepid. 9(2): 89-96. [See also p. 65 under “Entomological.” ]

& D. McFaruanp. 1977. Introduction to Howatharra Hill Reserve in the
Moresby Ranges near Geraldton, Western Australia. Printed by Geraldton News-
papers. [Available at cost from the author: P.O. Box 56, Northampton, Western
Australia 6535. |

Meapty, G. R. W. 1965. Weeds of Western Australia. Dept. of Agriculture, Perth.

ParuaM, J. W. 1964. Plants of the Fiji Islands. Govt. Press, Suva, Fiji.

SmirH, G. G. 1973. A guide to the coastal flora of South-Western Australia. Hand-
book No. 10, W. Aust. Naturalists’ Club, Perth.

TuHorNnE, R. F. 1968. Synopsis of a putatively phylogenetic classification of the
flowering plants. Aliso 6(4): 57-66.

WiiuiaMs, R. J. 1955. Atlas of Australian resources. “Vegetation regions’—color
map and leaflet. Dept. National Development, Canberra. 23 pp.

Wiis, J.C. 1973. A dictionary of the flowering plants and ferns, 8th ed. (revised
by H. K. Airy Shaw). Cambridge Univ. Press, London.

ENTOMOLOGICAL:

Common, I. F. B. 1966a. A new family of Bombycoidea (Lepid.) based on Car-
thaea saturnioides Walker from Western Australia. J. Ent. Soc. Queensland 5:
29-36.

1966b. Australian moths (revised ed.). Jacaranda Pocket Guides. Jaca-

randa Press, Brisbane.

1970. In The insects of Australia [Chapt. 36, pp. 765-866]. Melb. Univ.

Press, Melbourne.

1974. In The insects of Australia, Supplement [pp. 98-107, 133]. Melb.

Univ. Press, Melbourne.

1977. A new genus Myrascia for a group of Australian Oecophoridae

(Lepid. ) previously referred to Philobota Meyrick. J. Aust. Ent. Soc. 16: 87-109.

1979. The larva and pupa of Imma acosma (Turner) and I. vaticina

Meyrick (Lepidoptera: Immidae), and the taxonomic relationships of the family.

J. Aust. Entomol. Soc. 18: 33-38.

& T. E. Bextias. 1977. Regurgitation of host-plant oil from a foregut

diverticulum in the larvae of Myrascia megalocentra and M. bracteatella (Lepid.:

Oecophoridae). J. Aust. Ent. Soc. 16: 141-147.

& N. McFarianp. 1970. A new subfamily for Munychria Walker and
Gephyroneura Turner (Lepid.: Anthelidae) and the description of a new species
from Western Australia. J. Aust. Ent. Soc. 9: 11-22.

Forses, W. T. M. 1958. Caterpillars as botanists. Proc. 10th Int. Congr. Ent. 1:
313-317.

GoutprincH, G. M. 1929. Revision of the Australian Geometridae—Geometrinae
(Archaic genera). Proc. Linn. Soc. N.S.W. 54(4): 379-407 + plates XIV—XVI.

McFartanp, N. 1964. Notes on collecting, rearing, and preserving larvae of Macro-
lepidoptera. J. Lepid. Soc. 18(4): 201-210.

1965. Additional notes on rearing and preserving larvae of Macrolepidop-

tera. J. Lepid. Soc. 19(4): 233-236.

1970. “Moth Eggs!” Aust. Nat. Hist. (mag.): 346-352 (June, 1970). [A
semi-popular article with egg photos depicting 19 Australian spp. ]

———. 1972a. Notes on describing, measuring, preserving and photographing the
eggs of Lepidoptera. J. Res. Lepid. 10(3): 203-214.

SUPPLEMENT TO VOLUME 33 65

. 1972b. Egg photographs depicting 40 species of southern Australian moths.

J. Res. Lepid. 10(3): 215-247.

. 1973. Some observations on the eggs of moths and certain aspects of first

instar larval behavior. J. Res. Lepid. 12(4): 199-208.

. 1974. In memory of Eileen Victoria and James Otto Wilson. Aust. Ent.

Mag. 2(2): 40-42.

. 1975. Larval foodplant records for 106 species of North American moths.

J. Lepid. Soc. 29(2): 112-125.

1976. Hilltopping and defence behaviour in a diurnal agaristid moth. Aust.

Ent. Mag. 3(2): 25-29 [ref.—my As.5].

. 1977. See under “Botanical’—p. 64.

McGauran, J. 1951. The life history of the brown-tail moth, Pterolocera isogama
Turner ( Anthelidae). W. Aust. Nat. 3(2): 24-26.

Mitts, M. B. 1950. Observations on processionary caterpillars. W. Aust. Nat. 2(4):
84-87.

1951-52. Bag shelter caterpillars and their habits, Parts I & II. W. Aust.

Nat. 3(3): 61-67; 3(4); 84-92.

1954. Observations on the life history of the moth, Anthela xantharcha
(Meyrick) (Anthelidae). W. Aust. Nat. 4(4): 86-90 [ref—my An.12 and
An.18].

Muruura, A., Y. YaMAMmoro, & I. Hatrort. 1970. Early stages of Japanese moths
in colour, Vol. I. (2nd ed., rev. by S. Issiki.) Hoikusha Pub. Co., Osaka.

Ropinson, G. S. 1975. Macrolepidoptera of Fiji and Rotuma. E. W. Classey Ltd.,
England.

SHIELDs, O., J. F. EMMEL, & D. E. BREEDLOVE. 1970. Butterfly larval foodplant
records and a procedure for reporting foodplants. J. Res. Lepid. 8(1): 21-36.
Sincu, B. 1953. Immature stages of Indian Lepidoptera, No. 8—Geometridae.

Indian For. Rec. (Ent.) 8(7): 67-158 + i-ii, 10 pls. [Agathia larva and others].

TmLLyARD, R. J. 1926. The insects of Australia and New Zealand. Angus and
Robertson, Sydney. 560 pp.

Witson, J. O. 1972. A new species of Thalainodes (Lepid.: Geometridae—En-
nominae ) from Central Australia. Mem. National Mus. Victoria 33: 123-24 [ref.
—my G.180].

66 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

CoMMON NAMES OF FOODPLANTS IN THE INDEX THAT FOLLOWS:

Arum Lily—see Zantedeschia.
Ash—see Fraxinus.
Aster Family—see Composites.

Bacon and Eggs—see Pultenaea.

Beach Grass—see Spinifex.
Bindweed—see Convolwvulus.

Birch—see Betula.

Blackboy—see Xanthorrhoea.
Boneseed—see Chrysanthemoides.
Bracken—see Pteridium spp.

Bull Oak—see Casuarina.

Bur Medic or Bur Clover—see Medicago.

Cape Dandelion—see Arctotheca.

Cape “Ivy’—see Senecio mikanioides.

Capeweed—see Arctotheca.

Carrot, Native—see Daucus.

Champion Bay Poison Bush—see Gastrolobium.

“Cherry ’, Native—see Exocarpos cupressiformis.

Chickweed—see Stellaria.

Christmas Bush—see Bursaria.

“Cranberry, Native—see Astroloma humifusum.

Composites—see Arctotheca, Artemisia, Chrysanthemoides, Crassocephalum, Olearia,
Pterocaulon, Senecio, Solidago.

Cooktown Ironwood—see Erythrophleum.

Daisy Family—see Composites.
Devil’s Twine—see Cassytha spp.
Dock—see Rumex.
Dodder-Laurel—see Cassytha spp.
Double-gee—see Emex.

Evening Primrose—see Oenothera.

Fat Hen—see Chenopodium.

Ferns—see Nephrolepis; Pteridium spp.
Five-corner—see Averrhoa.

Fungus—see “fungus (unidentified )”; Psaliota.

Goldenrod—see Solidago.

Grape Family—see Cayratia, Cissus, Clematicissus, Vitis.
Grape, Native—see Clematicissus.

Grasses—see Poa, Spinifex; also see “grasses (unidentified )”.
Grass-tree—see Xanthorrhoea.

Gums—see Eucalyptus spp.

Heath Family (Epacrids )—see Astroloma, Leucopogon.
“Honeysuckle”, Native—see Banksia marginata.

[ronwood—see Erythrophleum.
Ivy—see Hedera; also Senecio mikanioides.

LLegumes—see Pea Family.
Lemon—see Citrus.

SUPPLEMENT TO VOLUME 33 67

Lignum—see Muehlenbeckia.
“Lilac”, Native—see Hardenbergia.
Loquat—see Eriobotrya.

Mallees—see Eucalyptus diversifolia, etc.
Mallows—see Alyogyne, Hibiscus, Lagunaria, Malwa.
Mint—see Mentha.

Mistletoes—see Amyema spp.

Morning Glory—see Convolvulus.

Moss—see Pottia.

Mushroom—see Psaliota.

Needlebushes—see Hakea spp.

Old Man’s Beard—see Clematis.
Oldman Saltbush—see Rhagodia.

Paperbark—see Melaleuca lanceolata.

Paterson’s Curse—see Echium.

Pea (edible )—see Pisum.

Pea Family—see Acacia, Bossiaea, Brachychiton, Brachysema, Cassia, Daviesia,
Dillwynia, Crotalaria, Erythrophleum, Gastrolobium, Genista, Hardenbergia, Hovea,
Jacksonia, Lotus, Medicago, Pisum, Platylobium, Podalyria, Pultenaea, Templetonia.

Peppermint Box—see Eucalyptus odorata.

Pepper Tree—see Schinus.

Pigweed—see Chenopodium.

Pimpernel—see Anagallis.

“Pine”, Native—see Callitris (Cupressaceae ).

Plum—see Prunus.

Potato Weed—see Heliotropium.

Primrose, Evening—see Oenothera.

Rattlepod—see Crotalaria.
Rhubarb, wild—see Rumex.
Ribwort—see Plantago.

Sagebrush—see Artemisia.

Saltbush, Oldman—see Rhagodia.

Salvation Jane—see Echium.

Sedges—see Gahnia; “sedges (unidentified )”’.
Sheoak—see Casuarina spp.

Silky Oak—see Grevillea robusta.
Smokebush—see Conospermum.
Star-fruit—see Averrhoa.

Stringybark—see Eucalyptus obliqua.

Tea Tree—see Leptospermum spp.
Thickhead—see Crassocephalum.
Three-cornered Jack—see Emex.
Tobacco, Tree—see Nicotiana.

Wattles—see Acacia spp.
Wireweed—see Cassytha; Polygonum.

Yakka—see Xanthorrhoea.

68 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
INDEX TO SCIENTIFIC NAMES OF FOODPLANTS!':

ACACIA (sp. unidentified )—36, 41, 48.
acuminata (W )—49.

anceps—60.
aneura—60.
armata—50.

* baileyana—33, 41.
brachybotrya—33.
calamifolia—29.
ericifolia (W )—49.
*iteaphylla—34.
leptostachya (Q)—47.
ligulata (W, in part )—15, 22, 23, 25, 27, 28, 36, 39, 49, 50, 52, 53.
mearnsii—3A4.
mollissima—34.
myrtifolia—21, 35.
oxyclada (W )—49.
pycnantha—15, 16, 21, 22, 23, 24, 26, 27, 28, 32, 33, 34, 35, 36, 39, 40, 46, 50,
pil oo; 60:
rivalis—35, 40.
rotundifolia—25.
salicina—27, 36, 39.
Psaligna (W )—49.
tetragonophylla (W )—49.
ulicina (W )—49.
victoriae—39.
ACANTHOCARPUS preissii (W )—61.
algae (unidentified )—18.
ALYOGYNE hakeifolia (W )—52.
ALYXIA buxifolia—22.
spicata (Q)—35.
AMYEMA melaleucae—14.
miquelii—14, 47.
*“ANAGALLIS arvensis—42.
* ARCTOTHECA calendula—13, 17, 27.
*ARTEMISIA californica (W)—28.
ASTROLOMA conostephioides—54.
humifusum—42.
*AVERRHOA carambola (Q)—389.

BAECKEA behrii (V)—21, 23, 26, 29, 43, 45.
BANKSIA attenuata (W )—56.

caleyi (W)—19.

marginata—16, 55.

menziesii (W )—56.

ornata—55.

sphaerocarpa, s.l. (W )—16, 19, 44.
BERTYA mitchellii—28, 34, 41.
*BETULA (sp. unidentified )—50, 57.

1 NOTE: In this foodplant index all species listed are South Australian except where other
states are indicated by abbreviations (in parentheses after plant names) as follows: (NS) = New
South Wales; (NT) = Northern Territory; (Q) = northeastern Queensland; (V) = Victoria;
(W) = Western Australia. An asterisk (*) before the name designates a naturalized or introduced
plant not native to the locality named in the preceding list.

See pp. 9-11 for a general discussion of many of the S. Aust. plants listed here, plus some other
i EcaMied spp. not in this index. Only plants associated with larval feeding records are indexed
1eTe,

SUPPLEMENT TO VOLUME 33 69

BEYERIA (sp. unidentified )—21.
leschenaultii var. latifolia—22, 34, 40.
opaca—21.

BOERHAVIA (sp. unidentified) (NT )—58.
chinensis (W )—56, 58.
diffusa—58.

BOSSIAEA biloba (W )—357.

*BRACHYCHITON populneum—53.

BRACHYSEMA aphyllum (W)—49.

BURSARIA spinosa—18, 22, 24, 26, 28.

CALLITRIS columellaris ssp. intratropica (Q)—52.
preissii—22, 28, 40.
rhomboidea ( V )—28.

CALOTHAMNUS homalophyllus (W )—59.
validus (W)—59.

CALYTRIX involucrata—43.
tetragona—21, 25, 43.

*CAMELLIA (sp. unidentified )—24.

CASSIA (sp. unidentified )—15, 33.
nemophila—15, 27, 28, 30, 33, 34, 60.
CASSYTHA (sp. unidentified )—14, 24, 37.
glabella—14, 21, 37.
pubescens—14, 21, 25, 29, 32, 37.
CASUARINA (sp. unidentified )—15, 48.
littoralis (Q)—48, 58, 61.
muelleriana—15, 18, 21, 24, 59.
paludosa var. robusta—41.
striata—16, 51.
stricta—15, 18, 22, 24, 48.
suberosa—see littoralis.
CAYRATIA ?clematidea (Q)—13.

*CENTAURIUM (sp. unidentified )—42.

CHAMAESYCE australis ssp. glaucescens—51.
coghlanii—51.
sharkdensis—51.
CHENOPODIACEAE (unidentified )—52.

*CHENOPODIUM (sp. unidentified )—42.
CHORETRUM glomeratum—50.

spicatum—16.

*CHRYSANTHEMOIDES monilifera—17, 21, 22, 26.
CISSUS opaca (Q)—13.

*CITRUS (sp. unidentified, probably lemon )—28.
CLEMATICISSUS angustissima (W )—58.
CLEMATIS microphylla—41.
CONOSPERMUM stoechadis (W )—44, 45.

*CONVOLVULUS (sp. unidentified )—57.

erubescens—51.

*COPROSMA baueri—38.

*CRASSOCEPHALUM crepidioides (Q; Fiji) —19, 63.
CRASSULA (sp. unidentified )—50.
CROTALARIA trifoliastrum (NT)—18.
CRYPTANDRBRA (sp. unidentified) (W )—23.

tomentosa—23, 25.
Cryptostemma—see Arctotheca.

70 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

DARWINIA micropetala—35, 59.
DAUCUS glochidiatus—16, 50.
DAVIESIA brevifolia—16, 32.
divaricata (W )—22.
hakeoides (W )—54.
DILLWYNIA hispida—25, 36.
DODONAEA bursariifolia—28, 37.
*multijuga (NS )—56.
triquetra (NS )—37.
viscosa—24, 26, 31, 46, 50.
DRYANDRA (sp. unidentified) (W )—19.
cirsioides (W )—19, 20, 55.
fraseri (W )—20, 44.
pteridifolia (W )—19.

*ECHIUM lycopsis—17, 18.

*E MEX australis (W )—58.

EPILOBIUM (sp. unidentified )—14.
EREMOPHILA freelingii—15.

*ERIOBOTRYA (sp. unidentified )—24.
ERYTHROPHLEUM chlorostachys (Q)—15.
EUCALYPTUS (sp. unidentified) (W, in part)—56, 59, 60, 62.

alba (Q)—38.
baxteri—57.
* cladocalyx—28.
diversifolia—31, 37.
?drepanophylla (Q)—39.
fasciculosa—38, 62.
*ficifolia—21, 31.
leucoxylon—29, 31, 32, 39, 45, 48, 57.
obliqua—38.
odorata—23, 27, 28, 29, 30, 31, 32, 33, 37, 38, 39, 42, 43, 45, 46, 47, 48, 49,
50, 54, 56, 57, 58, 59, 60, 61.
Ppolycarpa (Q)—38, 39, 40, 46, 47.
viminalis—31.
EUPHORBIA spp.—see Chamaesyce—51.
EUROSCHINUS falcata (Q)—329.
EXOCARPOS aphyllus—33, 53.
cupressiformis—18, 21, 22, 28, 34, 36, 48, 50.
sparteus (W)—33.
syrticolus—33.

FENZLIA obtusa (Q)—27, 37, 43, 47, 54.
ferns—29, 30, 51.
*“FRAXINUS (sp. unidentified )—17.
*FUCHSIA (sp. unidentified )—58.

fungus (unidentified )—53.

GAHNIA lanigera—16.
GASTROLOBIUM oxylobioides (W )—49.
*GENISTA maderensis—17, 26, 57.
grasses (unidentified )—13, 15, 16, 17, 52.
*GREVILLEA (sp. unidentified )—45, 55.
concinna (W )—19, 20.
ilicifolia—45.
lavandulacea—25.
pinaster (W )—25, 34, 44, 55.

SUPPLEMENT TO VOLUME 33

*robusta—45.
?trachytheca (W )—44, 46.
HAKEA (sp. unidentified )—45.
auriculata (W )—46.
francisciana—55.
lissocarpha (W )—46.
muelleriana—16, 45, 55.
rostrata—16, 25, 44, 45, 46, 55.
rugosa—21, 55.
trifurcata (W )—45, 55.
HALGANIA cyanea—51.
HALORAGIS heterophylla—41.
HARDENBERGIA violacea—28, 50.
*HEDERA (sp. unidentified )—28.
*HELIOTROPIUM europaeum—18.
HIBBERTIA (sp. unidentified )—42.
acicularis—9, 51.
exutiacies—51.
fasciculata—14.
sericea—51, 62.
stricta—A4l, 51, 62.
virgata—Al, 62.
HIBISCUS diversifolius (Q)—5l.
HOVEA longifolia—s7.
HYDROCOTYLE (sp. unidentified )—16, 50.

ISOPOGON ceratophyllus—44.

JACKSONIA (sp. unidentified ) (W )—49.
compressa (W )—55.
furcellata (W )—55.
Plehmannii (W )—55.
Psericea (W )—55.
spinosa (W )—55.

*LAGUNARIA patersonii—53.

*LANTANA (sp. unidentified )—50.
LEPIDOSPERMA (W)—17.
LEPTOSPERMUM (sp. unidentified )—26, 43.

coriaceum—43.
myrsinoides—21, 30, 32, 35, 38, 41, 43, 56, 59, 62.
LEUCOPOGON parviflorus—56.
LEUCOSYKE corymbulosa (Fiji)—63.
LHOTSKYA (sp. unidentified )—43.
lichens (unidentified )—18, 61.
liverworts (unidentified )—18.
*LOTUS scoparius (W)—28.
LYTHRUM hyssopifolia—42.

*MALUS (sp. unidentified )—17.
*MALVA (sp. unidentified )—13, 42.
*MEDICAGO polymorpha var. vulgaris—41, 42, 47, 52.
MELALEUCA gibbosa—35, 43, 46, 53, 54.
glomerata—33.
halmaturorum—33.
lanceolata (W, in part )—33, 62.
megacephala (W )—53, 56.

OL

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bo

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

oraria—33, 35, 43, 46, 54, 55.
radula (W )—56.
scabra, s.l. (W )—56.
uncinata (V, W )—23, 26, 55, 56.
*MENTHA (sp. unidentified )—42.
*MIRABILIS jalapa—s58.
moss—see Pottia.
MUEHLENBECKIA gunnii—28.
mushroom—see Psalliota.
MYOPORUM viscosum—50.
*MYOSOTIS—(sp. unidentified )—18.

*NEPHROLEPIS ?cordifolia (Q)—51.
*NICOTIANA glauca (W)—53.

*OENOTHERA hookeri—14.
OLEARIA axillaris (W, in part )—28, 54.
pannosa—54.
ramulosa—17, 21, 22, 26, 41.

*PELARGONIUM (sp. unidentified )—28, 53.
PIMELEA serpyllifolia—38.
stricta—38.
*PINUS (sp. unidentified )—17.
*PISUM (garden pea)—22.
*PLANTAGO lanceolata—17, 42, 47.
PLATYLOBIUM obtusangulum—16, 50.
*POA bulbosa—16.
*PODALYRIA (sp. unidentified )—57.
POLYGONUM (sp. unidentified )—42.
Paviculare—21, 28, 47.
POMADERRIS paniculosa—23.
POTTIA (sp. unidentified )—18.
*PRUNUS (sp. unidentified )—24.
*PSALLIOTA arvensis—53.
PTERIDIUM ?Paquilinum (Q)—380.
esculentum (Q, in part )—29, 30.
PTEROCAULON glandulosum (Q)—52.
sphacelatum (Q)—52.
PULTENAEA largiflorens var. latifolia—21, 22, 24, 25, 26, 28, 32, 34, 36, 41, 50, 54.

RHAGODIA parabolica—26, 52.
*RUMEX (sp. unidentified )—13.

*SCHINUS molle—28.
SCHOLTZIA Pparviflora (W )—20.
sedges (unidentified ) (W )—17.
SENECIO aff. lautus—19.

*mikanioides—19.

*SOLIDAGO (sp. unidentified )—41, 42.
SPINIFEX longifolius (W )—51.

*STELLARIA (sp. unidentified )—42.

TEMPLETONIA retusa (W )—57.
*VITIS (sp. unidentified )—14, 58.
XANTHORRHOEA semiplana—20.

*ZALUZIANSKYA diwvaricata (W )—27.
*ZANTEDESCHIA aethiopica-—57.

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EDITORIAL STAFF OF THE JOURNAL
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SUPPLEMENT TO VOLUME 33

ANNOTATED LIST OF LARVAL FOODPLANT RECORDS
FOR 280 SPECIES OF AUSTRALIAN MOTHS

NOEL MCFARLAND

SValume 33 1979 Naber 4

JOURNAL

_ LEPIDOPTERISTS’ SOCIETY

Published quarterly by THE LEPIDOPTERISTS’ SOCIETY

Publié par LA SOCIETE DES LEPIDOPTERISTES
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN
Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS

ial 8 April 1980

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Cover illustration: Third instar larva of Limenitis archippus Cramer (Nymphalidae)
preparing to enter winter diapause. The larva is resting on the lip of its hibernaculum
constructed from the basal portion of a chewed tubular willow leaf (Salix babylonica
Linnaeus) covered with silk. In the autumn such larvae begin facultative diapause in
response to decreasing day-length. Original drawing by Mr. George C. Ford, Jr., Graph-
ics Illustrator, Department of Biological Sciences, University of Maryland Baltimore
County, 5401 Wilkens Avenue, Catonsville, Maryland 21228.

JOURNAL OF

Tue LeEprimpoprerRists’ SOCIETY
Volume 33 1979 Number 4

Journal of the Lepidopterists’ Society
33(4), 1979, 209-215

FIVE NEW SPECIES OF THE TRIBE EUCOSMINI
(TORTRICIDAE)

ANDRE BLANCHARD
P.O. Box 20304, Houston, Texas 77025

ABSTRACT. Phaneta mayelisana, Phaneta verecundana, Eucosma atascosana,
Eucosma guttulana and Eucosma diabolana are described. Imagines, male and female
genitalia, and wing venations are represented.

Phaneta mayelisana A. Blanchard, new species
Dicey Da. LiL 1m

Head. Palpi exceeding head by about an eye diameter, white except on pale brown-
ish gray outer side of second segment, which bears on its underside a tuft of long white
scales exceeding and almost hiding downturned, white third segment. Front and vertex
white. Antennae simple, white; pubescence in male not exceeding the scales, still
shorter in female. Thorax: Patagia and mesonotum white, tegulae white, spotted with
pale brownish gray in their middle. Abdomen whitish.

Maculation (as in Fig. 1). Fasciae white; ground color of both wings a brownish
gray hue of variable saturation. Fringe of forewing white basally, peppered outwardly
_ with brownish gray. Fringe of hindwing white.

Venation (as in Fig. 17). This insect shares with Eucosma cataclystiana (Walker)
the unusual character that-veins M, and Cu, of the forewing fuse about midway be-
tween cell and termen. .

Hindwing: Rs and M, approximate toward base; M; and Cu, united.

Length of forewing. Males, 10.2-12.2 mm, mean = 11.1 mm; females, 11.5 mm
(single specimen).

Male genitalia (Fig. 6). Figured from slide A.B. 4322, paratype from Canadian,
Hemphill Co., Texas, 2.VI.70.

Female genitalia (Fig. 11). Slide A.B. 4319, paratype from Paducah, Cottle Co.,
Texas, 17.IV.68. The signa and the sclerotization of the ductus bursae are obscured
because the genitalia took more than the optimum amount of chlorazol black. Two
signa, the smaller signum more ventral. Ductus bursae with sclerotized band with a
iength about one and one-half times its diameter, separated from ostium by a short
membranous section. Ductus seminalis attached ventrally at middle of sclerotized
band. Lamella postvaginalis semicircular and well sclerotized.

Holotype. 6, Matador Wildlife Management Area near Paducah, Cottle Co., Texas,
17.1V.68, collected by A. & M. E. Blanchard, deposited in the National Museum of
Natural History (type No. 75817).

210 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fics. 1-5. Holotypes: 1, Phaneta mayelisana; 2, P. verecundana; 3, Eucosma atas-
cosana; 4, E. guttulana; 5, E. diabolana.

Paratypes. Same location, same date as the holotype, 6 3, 1 9. Gene Howe Wildlife
Management Area near Canadian, Hemphill Co., Texas, 13.1V.69, 6; 14.IV.69, 6;
29.V.70, 3; 2.V1.70, 2 6; all collected by A. & M. E. Blanchard.

Also in the National Museum are three specimens of this species: one collected by
F. H. Snow in Clark Co. Kansas, June, 1962 (No. 146); one from Denver, Colorado (No.
187), no date; one from Colorado (No. 523), no date. They are included here for dis-
tribution record, but I do not make them paratypes because they are in rather poor
condition.

Dr. J. F. Gates Clarke who has examined some of the paratypes comments: “In
pattern this is much like columbiana, but is a much larger insect.”

I take great pleasure in naming this pretty insect for my beloved wife who collected
it with me.

Phaneta verecundana A. Blanchard, new species
Figs. 2, 7, 12, 18

Head. Palpi projecting the length of the head beyond front, much compressed,
white with a faint grayish spot on outer side of second segment and a grayish shading

VOLUME 33, NUMBER 4 was

10

Fics. 6-10. Male genitalia: 6, Phaneta mayelisana; 7, P. verecundana; 8, Eucosma
atascosana; 9, E. guttulana; 10, E. diabolana.

toward end of tuft scales underneath second segment; third segment hidden by tuft.
Face and vertex white. Antennae white, shortly pubescent in male. Thorax: Patagia
white; tegulae white to faintly ochreous, mesonotum white. Forewing with arched
costa, termen oblique, concave between veins R, and Cu,

Maculation (as in Fig. 2). Forewing white with markings pale ochreous in males
but generally somewhat darker in females. Hindwing whitish to pale gray.

Venation (Fig. 18). Veins R, and M, of hindwing very closely approximate toward
base; veins M, and Cu, united.

Length of forewing. Males 7-9 mm, mean = 8 mm; females 7.3-8 mm, mean =
7.6 mm.

Male genitalia (Fig. 7). Slide A.B. 4340, paratype from Canadian, Texas, 13.VIII.71.

Female genitalia (Fig. 12). Slide A.B. 4341, paratype from Canadian, Texas,
28.V.70. Corpus bursae membranous, two signa present; ductus bursae with some scler-
_ otization around it near ostium.

Holotype. ¢, Gene Howe Wildlife Management Area near Canadian, Hemphill
Co., Texas, 15.VIII.71, collected by A. & M. E. Blanchard, deposited in the National
Museum of Natural History (No. 75818).

212 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

Fics. 11-16. Female genitalia: 11, Phaneta mayelisana; 12, P. verecundana; 13,
Eucosma atascosana; 14, E. guttulana; 15, E. diabolana; 16, E. graziella.

Paratypes. Same location as holotype, 28.V.70, 6, 4 2; 13.VIII.71, 4 3. Matador
Wildlife Management Area, near Paducah, Cottle Co., Texas 4.VI.70, 6; all collected
by A. & M. E. Blanchard. Dr. J. F. Gates Clarke who examined two of my dissected
paratypes commented as follows: ““Very similar to indagatricana, but in (verecundana)
the costal strigulae of the forewing are confined to the outer half of the costa, in in-
dagatricana they go nearly to the base of the costa. The genitalia also differ: in the
male the neck of the harpe (valve) is much narrower and the excavation of the ventral
edge of the harpe is much deeper in (verecundana) than in indagatricana; also in the
female of (verecundana) the postvaginalis is not sclerotized but is in indagatricana.”

Eucosma atascosana A. Blanchard, new species
Figs. 3, 8, 13, 19

Head light ochreous; palpi exceeding front by an eye diameter; tuft on underside of
second segment externally ochreous, slightly compressed, loosely scaled, concealing

VOLUME 33, NUMBER 4 INS

Fics. 17-21. Venations: 17, Phaneta mayelisana; 18, P. verecundana; 19, Eucosma
atascosana; 20, E. guttulana; 21, E. diabolana.

third segment. Antennae slightly compressed laterally, finely ciliate in male, light
ochreous. Thorax: Tegulae and anteromedial mesonotum ochreous brown; posterior
tuft white. Abdomen: whitish ochreous.

Maculation (as in Fig. 3). Forewing: rich yellowish brown, a little paler in the fold,
with silvery white spots showing negligible variation in all specimens before me. All
white spots, except the elongate one along the dorsum and the one near the costal fold,
surrounded by line of dark brown scales; fringe pale ochreous with a darker line near
base; male costal fold extending to about one fourth of the costa. Hindwing: pale
ochreous with concolorous fringe.

Length of forewing. Males 11.5-13.0, average 12.5 mm; females 11.0-14.0, average
12.0 mm.

Venation (Fig. 19).

_ Male genitalia (Fig. 8). Slide A.B. 4489, paratype from Laguna Atascosa, Cameron
Mos Lexas: 22.X1.73.

Female genitalia (Fig. 13). Slide A.B. 4459, paratype from Laguna Atascosa, Cam-
eron Co., Texas, 22.X.73. Corpus bursae and ductus bursae membranous; two large
signa; lamella postvaginalis about one and one half times as long as broad, lightly scler-
otized, with long setae.

Holotype. ¢, Laguna Atascosa National Wildlife Refuge, Cameron Co., Texas,
22.X1.73, genitalia on slide A.B. 4313, deposited in the National Museum of Natural
History (No. 75821); collected by A. & M. E. Blanchard.

Paratypes. Welder Wildlife Refuge near Sinton, San Patricio Co., Texas, 28.X.64,
6; 12.X1.65, 6; 13.X1.65, 2. Voshell Wildlife Management Area, near Brownsville,
Cameron Co., Texas, 12.X1.68, ¢; 5.XI.69, 6 2; 9.XI.69, 2; 26.X.70, ¢. Laguna Atascosa
Wildlife Refuge, Cameron Co., Texas, 19.X1.73, 6; 22.X1.73, 4 6, 7 2; collected by A.
& M. E. Blanchard.

This insect is close to Eucosma sandiego Kearfott, as shown by their male genitalia
and maculation. Dr. J. F. Gates Clarke who compared some of my specimens to the
sandiego specimens in the National Museum commented: “Generally the spots of your
species are larger and more rounded than in sandiego and the three subterminal spots
of your species are larger and more distinct than those in sandiego. The neck of the
harpe of your species is narrower than that of sandiego.”

214 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Eucosma guttulana A. Blanchard, new species
Figs. 4, 9, 14, 20

Head. Palpi exceeding front by half eye diameter; second segment white medially,
whitish above; external side and shaggy brush of long scales on underside tawny; third
segment smoothly scaled, downturned, tawny. Front and vertex white, spotted with
tawny. Antennae simple, tawny; pubescence in male barely exceeding the scales.
Thorax: mesonotum tawny with white anterior band and white posterior tuft. Tegulae
tawny with white tip. Abdomen pale ochreous above, paler beneath.

Maculation (as in Fig. 4). Forewing: ground color varying from ochreous to tawny
or even dark brown; spots white, except the three larger faintly ochreous ones forming
an ill-defined ocelloid patch. Fringe white, basally speckled with ground color. Hindw-
ing: slightly paler than ground color of forewing; fringe whitish, basally darker.

Venation (Fig. 20). Hindwing veins R, and M, approximate toward base; veins M,
and Cu, fused almost to termen.

Length of forewing. Males 7.5-12.0, mean = 9.5 mm; females 9.0-11.0, mean =
10.3 mm.

Male genitalia (Fig. 9). Slide A.B. 3597, paratype from Padre Island, Nueces Co.,
Texas, 9.1X.74.

Female genitalia (Fig. 14). Slide A.B. 4460, paratype from South Padre Island,
Cameron Co., Texas, 30.III.78. Papillae anales large, with a blunt ventral process turn-
ing caudodorsally. Lamella antevaginalis heavily sclerotized; lamella postvaginalis
broader, crescent shaped, weakly sclerotized; ductus bursae slightly sclerotized be-
tween junction with ductus seminalis and ostium, but not immediately cephalad of
ostium. Corpus bursae with a wide slightly sclerotized medial band that includes ven-
tral and dorsal signa.

Holotype. 6, Padre Island National Seashore, Kleberg Co., Texas, 19.VII.76, col-
lected by A. & M. E. Blanchard, deposited in the National Museum of Natural History
(No. 75819).

Paratypes. All from Texas: Engeling Wildlife Management Area near Tennessee
Colony, Anderson Co., 39.1V.66, 2 3; 6.1X.66, 6. Camp Strake near Conroe, Montgom-
ery Co., 27.1V.67, 3; 22.1V.69, 6. Matador Wildlife Management Area, near Paducah,
Cottle Co., 8.VIII.68, 6 6,3 2. Gene Howe Wildlife Management Area, near Canadian,
Hemphill Co., 28.V.70, 3. Welder Wildlife Refuge, near Sinton, San Patricio Co.,
30.VI.75, 3, ¢. North Padre Island, Nueces Co., 9.1X.74, 5 6, 9; 19.1X.74, 2 9;
12,111.75, 6: 30.1X.75, 2.3, 2; 17. VINL76). 2 So: 19. VILW7, 4 Se 21 Vile om eons
3, 2 2. Padre Island National Seashore, Kleberg Co., 2.X.75, 6, 2; 17.V.76, 2 6, 2
2; 19.V.76, 2; 19.VII.76, ¢. South Padre Island, Cameron Co., 1.111.78, 6; 30.11.78,
21 6,3 2. Attwater Prairie Chicken National Wildlife Refuge near Eagle Lake, Colo-
rado Co., 27.IV.78, 2 3; collected by A. & M. E. Blanchard.

Remarks. This species is closely related to Eucosma robinsonana Grote as shown
by the genitalia of both sexes and the wing venation, but the maculation is very dif-
ferent and it is a much bigger insect.

I have this insect only from eastern and southern Texas and from the Panhandle of
Texas: none from the wide intervening territory. The specimens from the Panhandle
are generally lighter in color (ochreous instead of tawny or brown) than those from the
East and South, but this appears to be no more than a color variation.

Eucosma diabolana A. Blanchard, new species
Figs. 5, 10, 15, 21

Head. Palpi exceeding front by half an eye diameter; second segment white an-
teriroly and medially, outer side pale brownish; underside with tuft of very long, dark
brownish scales greatly exceeding the smoothly scaled, downturned, half hidden third
segment. Front and vertex whitish. Antennae fasciculate in male, shortly pubescent
in female. Thorax: mesonotum and tegulae ochreous.

Maculation (as in Fig. 5). Forewing: from where the background is palest, near

VOLUME 33, NUMBER 4 215

apex, to the dark fasciae, the color is of about the same hue, varying only in saturation;
that is from a very pale ochreous near apex to a rich brown with an orange tinge in the
two large fasciae and near base along costa. Most wing scales and all the fringe scales
are white tipped. Hindwing: concolorous with the parts of the forewing with average
saturation only a little grayer.

Venation (Fig. 21). Hindwing: R, and M, approximate toward base, M, connate
with stalk of M; and Cu.

Length of forewing. Males 10.3-18.0, mean = 13.6 mm; females (three speci-
mens): 12.0, 12.7, 13.4 mm.

Male genitalia (Fig. 10). Slide A.B. 1233, paratype from Mt. Locke, Davis Mts.,
26.1I1.68.

Female genitalia (Fig. 15). Slide A.B. 4443, paratype from Sierra Diablo, 20.V.68.
Lamella antevaginalis a narrow sclerotized lip; lamella postvaginalis subquadrangular
with setae; ductus bursae membranous with narrow constriction near ostium. Corpus
bursae membranous with two minute signa.

Holotype. 6, Sierra Diablo Wildlife Management Area, 6,000 ft, Culberson Co.,
Texas, 31.11.70, collected by A. & M. E. Blanchard, deposited in the National Museum
of Natural History (No. 75820).

Paratypes. Davis Mts., Mt. Locke, 6,500 ft, Jeff Davis Co., Texas, 26.III.68, 6;
Sierra Diablo Wildlife Management Area, 6,000 ft, Culberson Co., Texas, 20.V.68, 5
pee ie Onor ol IL 70,6 6, 2: 3.1V.10,.6;27.V.73, 8 6, 2; 29.V.73, 2 36; 30.V.73,
2 5, collected by A. & M. E. Blanchard.

Remarks. “The smaller specimens (of E. diabolana) remind one of mirosignata
Heinrich, but your species is distinct and presumably undescribed” (Dr. J. F. Gates
Clarke, in litt.). The genitalia, male as well as female, are also very different.

Eucosma graziella A. Blanchard
Fig. 16

Remarks. This species was previously described (Blanchard, 1968) but the female
genitalia had not been studied. Fig. 16 is drawn from slide A.B. 4446, the genitalia of
a female taken in the Chihuahua Desert, near Nugent Mt. at Big Bend National Park,
Texas, 3.X.67. The ventral and lateral parts of the sterigma loosely surround the small
ostial chamber; its dorsal part extends caudad as a subquadrate lamella postvaginalis.
There is some slight sclerotization of the corpus bursae mediodorsally near the dorsal
signum.

ACKNOWLEDGMENT

I am deeply grateful to Dr. J. F. Gates Clarke for examining criti-
cally much of my material and comparing it with the material in the
National Museum. Without his unstinted help this article would not
have been possible. I also want to thank Mr. Fletcher of the BM(NH)

for helping me in the same manner.

LITERATURE CITED

BLANCHARD, A., 1968. New moths from Texas (Noctuidae, Tortricidae). J. Lepid. Soc.
222 VA3.

Journal of the Lepidopterists’ Society
33(4), 1979, 216-231

HOW TO MAKE REGIONAL LISTS OF BUTTERFLIES:
SOME THOUGHTS!

HARRY K. CLENCH?
Carnegie Museum of Natural History, Pittsburgh, Pennsylvania 15213

ABSTRACT. Procedures are described for making two types of regional lists of
butterflies: the state or provincial list, and the “local study” (an intensive, long-term
investigation of a small area). The need for such lists, problems in making them, and
some of the expectable results, are examined. A logarithmic scale for describing pop-
ulation sizes is given, as is a procedure for estimating total number of species in a local
area.

STATE AND PROVINCIAL LISTS

State or provincial lists are important sources of regional informa-
tion about a variety of aspects of butterflies, the particular aspects
being up to the writer. At the least such lists should include distri-
bution records, and they may also provide data on number and timing
of broods, habitat choice, rarity, phenotypic and geographic variation,
and so on.

As I pointed out many years ago (Clench, 1949), these lists are
invaluable to taxonomists, zoogeographers, ecologists, and to re-
searchers concerned with many other types of problems as well. They
are also useful to both resident and visiting collectors, showing when
and where to look for particular species. An often little-appreciated
value of such lists is in their “negative information” content: an omis-
sion can inform the collector of needed data—a still unknown food-
plant, an unguessed fall brood, as well as the more striking unrecord-
ed species—and thereby encourage him to publish any such newly
acquired information.

Methods

It is useful, in gathering and storing data, to keep two loose-leaf
notebooks: a species book and a county book.

The species book is the main list and comprises a separate sheet
for each species, more eventually if needed. On each species sheet
enter the full data for all records of that species that you acquire.
Enter the records as received, in no particular order. I always put the

' At the Lepidopterists’ Society meeting in Louisville in 1978 I gave a paper on this subject, and several of those
attending suggested that the information be made more permanently available. I here comply with that suggestion,
amplifying and extending the original paper in a few places.

This paper considers two kinds of regional lists: state lists or their equivalent; and what I call “local studies’ —
long-term investigations of small areas. Each requires a different approach, and each will be treated separately.

* Editor's note: This paper is published posthumously following Mr. Clench’s sudden and untimely death in April
1979. A future issue of the Journal will be published as a memorial to commemorate his many contributions to the
study of Lepidoptera, and to the Lepidopterists’ Society.

VOLUME 33, NUMBER 4 PAALTE

county name to the left in a column of its own, with the remaining
data in full to the right. The county column thus may be scanned
rapidly for particular records. Data entered on this sheet should com-
prise full locality data (including, when appropriate, elevation and
mountain range), date, source, as well as any other information asso-
ciated with the record. Do not omit the source. One of Murphy’s Laws
is that the record whose source you neglected to note will later be-
come critical.

It is also helpful to add a distribution map for each species. Such
a map, even if you only tick off the counties recorded (instead of
spotting all records exactly), will give you a good general picture of
the range of the species in your state, and the areas from which more
information is needed.

The county book contains a sheet for each county, parish, or equiv-
alent subdivision. Simply enter the names of all species recorded
from that county (referring to the species book for the full data). An
easy way to simplify the procedure is to type up a one page list of all
species known or strongly suspected to occur in the whole state. With
elite type, single spaced, and using three columns, you can get 150
species or more on one side of an 8% x 11” sheet. Add a blank line
at top right. Reproduce this sheet—by quantity-photocopying or mim-
eograph—to the number of counties in your state (get some addi-
tional copies: they are always useful). For each county, take one of
these sheets and write the county name on the blank line. Then sim-
ply check off the species you have recorded from that county. On the
reverse you can note sources of particular importance for that county:
published accounts, resident collectors, etc.

In the far west, where counties are particularly large, it may be
advisable to subdivide them into more useful smaller units for data
recording, in whatever arbitrary way is most appropriate to your
needs, and most easily and accurately described to others. The sub-
division preferably should be by linear, objective boundaries, such as
a river. Be careful using highways, however: they can be rerouted or
renumbered and cause confusion in later years.

Recording your data in these two books allows you to keep records
with a minimum of time, and a maximum of utility. You have instant
access to what you know about either a particular species, or a par-
ticular area.

Sources of Preliminary Data

Earlier lists of one kind or another are already published for many
_ states or provinces, or for parts of them. Such lists have often appeared
in obscure journals of limited distribution, and learning about them

218 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

can be a problem. An outstandingly useful source is Field, dos Passos,
and Masters, 1974. This bibliography, as its authors note, is not nec-
essarily complete, so you should do some extra searching. In addition
to earlier regional lists, monographs of various genera and other taxa
often contain many locality records.

Collections. (a) Large museums have at least some material from
almost every state and province. (b) Small museums, state colleges
and universities, state museums, nature centers, etc., often have local
and/or student collections. These may prove to be a gold mine of early
or otherwise unusual records, but watch for poor data. (c) Private
collections and collectors are probably your richest source of infor-
mation outside your own efforts. The local collector has often been
active for many years and may have exceptionally complete data on
the species in his area: identities, broods, timing, larval foodplants,
and so on, information of the utmost value and not duplicated else-
where. I have found such collectors to be extremely helpful and in-
variably more than willing to share their hard-won knowledge.

Problems. (a) Misidentifications. With experience you soon learn
which are the problem children: among them Erynnis, Hesperia, and
other hesperiids; Euphydryas, Speyeria, some lycaenids, Calephelis.
You may want to borrow specimens to confirm an identification, or
send your own to a specialist. In any event, do not hesitate to query
or even omit a dubious record, or you can assign it to the “hypothetical
list” (see below). “Thorybes sp.” is much preferable to “Thorybes
bathyllus” which is actually Thorybes pylades. The famous early Cu-
ban naturalist, Felipe Poey, has a wise saying: Mds vale ignorancia
que error (Ignorance is worth more than error).

(b) Stick to what you know. Do not assume any species—even Pieris
rapae—to be anywhere. Even common species may have gaps in their
ranges, and this would be important and interesting information. If
you find what seems to be a gap in the range of a species that
“shouldn’t” have such a gap, go check it out. It may be real.

Procedure in Fieldwork

Ideally your fieldwork should be conducted in two concurrent phas-
es: (1) an intensive, prolonged study of a small, easily accessible area
(see under “local studies” below); and (2) field trips to farther places
within your state or province.

The principal goal of the latter is to add as many distributional data
as possible, expressed (or summarized) in terms of county records.
Your aim is to acquire as much information as you can in the time at
your disposal. Initial efforts should be directed at simply adding as
many county records as possible. Pick the season when the most

VOLUME 33, NUMBER 4 219

species are flying (often June and July, but not always or everywhere),
and visit as many counties as time and funds permit. A fairly respect-
able list of species and county records eventually will be amassed,
and your attention may then gradually shift to intensified efforts at
more specific goals: a county still poorly represented; a species that
ought to be present but of which you have few records or none. The
search for such a species is more efficient if you first familiarize your-
self with available information about its habits, larval food, flight pe-
riods, and so on. Remember, some species fly only in particular sea-
sons, especially the spring, so seasonal collecting should be added to
your field exploration program.

Although you can think of the task as one of accumulating county
records, always remember that that is not the real goal but merely a
simplified accounting procedure, useful for record-keeping and sta-
tistical treatment. With that in mind, it may help to discuss some
possible patterns you may observe.

Every county has some upper limit to the number of species in it.
In a reasonably diversified state some counties will have relatively
many species, others fewer. If the counties are more or less uniform
in size and range of habitats, then the number of species in them
should be distributed in a Gaussian or normal curve, as in Fig. 1B.

Before you have begun to accumulate records, all counties in the
state have precisely zero species known from them: they would all
be ranged in a single bar at the extreme left of such a curve. As you
acquire records the counties begin to move up the graph to the right.
A single visit to a hitherto uncollected county may result in a dozen
or so species records, shifting that county up one class interval to the
right. With more visits the number will gradually increase. In Penn-
sylvania, a state I know well, a reasonably well-worked county—sev-
eral visits in different seasons—will have records of some 20-60
species. To increase the number beyond that point takes far more
effort, generally possible only by residence or intensive local col-
lecting.

After some years of work, both on your part and on the part of earlier
workers as well as collaborators in various parts of the state, the dis-
tribution of counties according to number of species will look some-
thing like Fig. 1A, which shows the current state of knowledge of
Pennsylvania butterflies. Note that the curve at this time is essentially
three-humped: (a) a group of 25 counties (37% of the 67 counties in
the state) with fewer than 20 records each, representing those either
collected in briefly or not at all; (b) a group of 28 counties (42%) with
between 20 and 60 records each, representing those visited a number
of times and in different seasons; and (c) a group of 12 counties (18%)

220 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

30 30

20

20

10

NO. OF COUNTIES
—_
e)

NO OF COUNTIES

Oo 20 40 60 80 100 0) 20 40 60 80 100
RECORDS PER COUNTY RECORDS PER COUNTY

Fic. 1. A. Present state of knowledge of Pennsylvania butterflies. Note the three
humps in the curve: a group of counties with fewer than 20 records each; a group with
between 20 and 60 records each; and a group with 60 or more records each. B. If all
Pennsylvania counties were completely known, the curve would probably look essen-
tially like this.

with 60 or more recorded species each. These last are counties with
a long history of collecting (Allegheny and Philadelphia counties); or
with collectors long resident (Lancaster Co. [George Ehle], or Tioga
Co. [George Patterson]); or in which especially intensive, long-term
collecting has been done (Westmoreland Co., where Carnegie Mu-
seum has a field research station). The distribution of collecting in-
tensity in Pennsylvania as measured by the number of species known
from each county, is shown in Fig. 2.

If the curve in Fig. 1B is summed (midpoint of each class interval
times number of counties in the class interval, and these totalled), we
have a theoretical maximum possible number of county records for
Pennsylvania of about 5,715. The total number now actually known
is 2,215, or 39% of those possible. Although this seems like a small
number, it is the result of many thousands of hours spent in the field
by collectors over more than a century. It shows dramatically how
difficult it is to get truly thorough knowledge about even one state.
I should add, however, that because several areas have been inten-
sively studied, about 145 species are now known from Pennsylvania,

VOLUME 33, NUMBER 4 Dahl

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Fic. 2. Distribution of collecting intensity by counties in Pennsylvania. Counties
with no shading have 20 or fewer records each; those with light shading have between
20-60 records each; and those with dark shading 60 records or more each.

and this total is not likely to be much increased in the future: perhaps
by no more than about 10.

Special Matters

Wherever you may be, you can help other lepidopterists. Perhaps
a taxonomist revising a group needs material or information from your
state; or a geneticist may want to find a colony of a particular species
in your area, or someone working on Monarch migration might need
local dates of immigration or emigration. For all such people the spe-
cial knowledge you have acquired could be of great value. And do
not forget reciprocity: his (or her) special knowledge may be of par-
ticular value to you someday as well.

Be aware of research that already has been done and that is relevant
to your area, and make an effort to extend the results. One frequent
problem is the nature of the boundary between two wide-ranging
subspecies. If that boundary passes through your state, and the ma-
terial you have acquired is adequate, you are uniquely situated to add
precision to what is known.

You may discover new problems. No general rule applies here.
Keep an inquiring mind, and be alert for things that “don’t fit.” Per-
haps you know of a local population with a different flight period than

iplepd JOURNAL OF THE LEPIDOPTERISTS SOCIETY

the species has elsewhere, or with a higher-than-usual frequency of
some dimorphic form. Or you might have noticed a persistent and
anomalous absence of some species from areas where it should be
found. A new species turning up in your area might represent a range
expansion, something we know little about.

The Goal

The objective in preparing a state list is the accurate record of con-
ditions in a particular place—your state—at a particular time or times,
and in as much detail as you are capable of.

Accuracy is vital. Rumored occurrences and sight records are ad-
missible only if they are clearly so reported. Misidentifications, ide-
ally, should never occur. Some few are bound to, but make every
effort to eliminate them completely: submit doubtful specimens to a
specialist; record any uncertain identifications as such. Be meticulous
with specimen data, and ultracareful in recording the information.

Data on brood numbers and timing, on larval foodplants, and other
attributes, should be clearly identified as to geographic source. If you
have no local data it is proper and even wise to copy from the liter-
ature (because it will provide clues for future users of your list), but
you should clearly indicate that the information was not locally ob-
tained.

Be aware of old records not recently duplicated; of species common
now but not mentioned by “the old boys.” Such things may seem
unimportant but could fit in with data from other areas to demonstrate
a pattern. A recent instance of this began with the account of Nathalis
iole by Kimball (1965). He remarked on the absence of early records
from the state and concluded that it might have established itself in
Florida relatively recently. I became interested, checked two large
museum collections and other early literature and concluded that iole
indeed previously was absent from Florida, that it reached the state
in 1913 or shortly before, and spread north from its probable original
landfall in the Keys (Clench, 1976).

Miscellaneous

A state list may invite a zoogeographic or ecological analysis, or
other derivative study. If your interests lie in these directions, so
much the better: a list can only be improved by such work. But these
“extras” are not necessary. What is necessary is a careful, accurate
compilation of reliable information. If you provide this, then your list
will be a valued and respected contribution to our science, useful and
used for a long time to come.

VOLUME 33, NUMBER 4 223

The “hypothetical list.” Ornithologists are familiar with this term;
lepidopterists are not. In your published list itself, include only
species with established reliable records. Save the rest for the “‘hy-
pothetical list’ at the end. It is the perfect place for the doubtful
species, the “possibles,” those previously reported in error. Anything
dubious can go in. This device allows you to avoid the difficult de-
cision of what to include in, and what to exclude from, the main list.
Amplify and discuss the entries as you will. The “hypothetical list”
is a wonderful place for the reader to browse when your paper is
eventually published.

The work of assembling a state list is never done. Information al-
ways remains to be learned, and always will. The trick is knowing
when you have reached the point when you can properly say, “This
is now worth publishing.” Eventually the time comes when you have
a reasonable picture of the butterflies of your state, embodying a sat-
isfying quantity of new data. If at this point the influx of new infor-
mation slows, then the time clearly has come to put it all together and
get it into print.

As the foregoing should have intimated, preparing a state list is not
a simple task. It requires experience, knowledge, and judgement, as
well as diligence. It is not really a job for a beginner. If you are a
beginner, however, and you really want to undertake such a task, then
you should seek as much advice and help as you can from those with
more experience. If you do that, then there is no reason not to produce
an excellent and valuable piece of work.

LOCAL STUDIES

A local study, as I use the term here, is an intensive, long-term
investigation of a small area: perhaps one or two thousand acres, about
as much as can be covered reasonably well on foot in a single day.

Studies of this kind have been undertaken so seldom that wherever
you choose to do so you will be rewarded with significant data, well
worth publishing. Because it requires no extensive collections and no
large reference library this kind of study is particularly suited to the
serious lepidopterist in a rural area. Bear in mind that much of the
information you acquire, even though it seems of little interest in
itself, will gain greatly in value when added to similar information
from elsewhere. Among the most important eventual results of such
work will be establishing geographic patterns of variation in brood
numbers and timing, larval foodplant choice, abundance levels, and
other things about which we are still totally, or almost totally, igno-
rant.

224 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

Define or enunciate what you want to do at the outset, and give it
careful thought. One important aspect of a local study is time varia-
tion, for which you need data extending over as long a time as pos-
sible. The earlier you start keeping records, the longer your time-span
of useful information.

Do not make your goals too many or too complicated or you will
defeat your own purposes. Know about how much time you will have
for the work and plan accordingly. After your first month or two, re-
view your program and your goals. Perhaps you have bitten off more
than you can chew; or perhaps you could easily do more than you
first thought.

The place: what to look for. Frequent visits are important. The
nearer your chosen spot is to your home, the more often you can visit
it.

The study area you choose should be reasonably representative of
your region; with enough habitat diversity this should be so. In a
pinch, however, almost any area will do: even a vacant lot in town,
if you cannot get to anything better. Nevertheless, in a day’s survey
you can record useful data on a large number of species, so (within
reason) the more diverse the area the better.

Your study area should have—and have had—no pesticide spraying.
It should be free from any abrupt, major change in land use: you do
not want the place bulldozed for a shopping center a year after you
begin! And it should be as free of people as possible. When you are
hard at work in your area you need to concentrate, to be free to follow
this or that butterfly, to observe what it does. People, even the best
intentioned, interfere with this work.

If you are lucky, a weather station (government or private) may be
located in or near your study area, and you can arrange for copies of
the data. Failing this, then look for a weather station as near as pos-
sible, and in as similar a habitat as possible, and record what you can
in your own area on your own visits. This is not as satisfactory, but
some idea of the regional climate is necessary.

State or local parks, perhaps even a national park, if they fit other
requirements of your study, may include suitable sites. If permits are
required, you must get them. In any event you should get permission.
The personnel of the park may even be able to help you locate the
best place for your study, if you explain your needs and aims.

Universities and other institutions often have their own study areas,
designed for investigations of just this kind. If so, they are ideal: they
are stable in land use, free of insecticides, often have climatic records,
and frequently have background data on vegetation, land use history,
maps of habitat types, and so on. They may also have a policy of

VOLUME 33, NUMBER 4 225

limiting the area to their own personnel. Sometimes, however, they
are only too happy to let you use the area, if you ask first, explain your
aims, and don’t mistreat the area.

Private lands often have ideal places for such a project. Again, ask
the owner and get his permission first. In recent years the land owner
has gotten much more hard-nosed about strangers on his land. He or
his neighbors may have been victimized by careless or malicious
campers, wanton vandals, drunken or stupid hunters, and who knows
what else: his attitudes can hardly be faulted. In most cases, however,
if you can satisfy him that you are serious, and that you will not tram-
ple his crops or leave trash behind you, he will willingly let you use
his property.

In using any land ordinary courtesy is necessary. Do not litter; do
not walk on crop plants; do not leave gates open if they were closed,
and vice versa, follow paths as much as possible. If the area is already
in use by other researchers, familiarize yourself with what they are
doing and be careful not to interfere with their projects. Respect fully
any ground rules that may be in force. Regardless of who owns the
land, it is good public relations to keep them informed, if only in a
general way, of your progress. If you find a rarity, or make an impor-
tant or unusual discovery, tell them. Land owners or managers like
to hear that their place is “special.” And when you publish, acknowl-
edge them by name, and give them a reprint of your paper.

Methods

I assume that you have chosen an area, and that you will be making
periodic visits, perhaps once a week or so. As in state lists, I keep two
books of records:

(1) Log book. This need not be loose-leaf, as it is strictly chrono-
logical: on each visit enter date, time you begin fieldwork, and time
you finish. Record weather data (temperature, cloud cover, wind; and
any important changes during your visit). List the species you take or
observe in the area, where you see them, how common they are, their
condition: these three matters are discussed more fully below. Record
any special observations, such as territorial behavior (time of day,
territorial activity, size of territories, and so on), predator attacks (de-
tails), mating or courtship behavior (nature of activity, time, sex of the
flying partner in copulating pairs, and so on), unusual numbers, ovi-
position records, flowers serving as adult food, etc.

(2) Species book. This should be loose-leaf, to allow additions. A
separate sheet (or more) for each species, entering dates, particular
places, and other information from your log. It will be repetitive, cer-
tainly, but this accumulation of data is the core of your whole project.

226 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

You will need to accumulate a voucher collection, either keeping
it yourself, or giving it to some inititution. It should comprise reason-
able series of each species, and of each brood, and of any problem
groups or unusual specimen for which documentation or later study
may be necessary. After this, collect as little as possible: the job is
primarily one of observation and recording. With a little familiarity,
most species can be identified without capture. Certain groups, such
as Polygonia, Erynnis, or some of the smaller hesperiids, may need
to be captured for reliable identification. Many can then be released,
but some must be kept for more careful identification later. Be alert
for the rare species that in the field looks much like some common
species.

The following procedures can improve the accuracy and detail of
your observations, and hence their later utility. Your study area can
be divided into a number of “microlocalities.” It is important to be
able to specify these, in order to localize observations on habitat
choice, colony locations, and so on. Two basic methods are used:

(1) A grid system, in which the whole area is marked off by a rect-
angular grid. In one type of grid the squares are identified, “B-6,” for
example, referring to the square in row B, column 6. In the other type
of grid the lines are identified, so that a square may be specified by
the intersection of some standard corner, or a point identified by a
fractional designation such as “5.6 E, 6.1 N,” meaning 0.6 unit east
of line 5 and 0.1 unit north of line 6. A grid system can be particularly
precise, but it requires more than just drawing the lines on a map:
you must have some means of identifying them on the ground, too,
or they are of little use. Survey markers along trails or at grid line
intersections is one method. Generally, if the area you use does not
already have a grid system, it is too involved and expensive to set one
up.

(2) Place names. Sometimes an area under study will have them, or
some of them already. If so, use them. If they do not exist, then you
will need to make them up. Do so with thought. It is tempting to use
such terms as “Idalia Meadow” or “Hypaurotis Scrub,” but such
names may sometimes cause confusion in your notes. The same may
be true of botanical adjectives. Other kinds of names can be more
practical, sometimes even silly ones. One of our places at Powdermill
is called “Elephant Walk.” ’'m not sure why, but it is certainly easy.
to remember. “Typha-Acoris Marsh” is there, too, and poses no prob-
lem, for that combination of plants occurs at Powdermill in only one
place. However you do it, keep a record on a map of the locations,
and document them with photographs.

Recording the condition of the specimens you see is extremely use-

VOLUME 33, NUMBER 4

bo
bo
—l

ful, particularly when visits to the area are at weekly or greater inter-
vals, because they give a valuable clue to how long the species has
been flying. I use a series of five lower case letters: a, perfect and
unblemished; b, showing slight wear; c, definitely worn; d, ex-
tremely worn; and e, a complete rag, the wings so rubbed that iden-
tification may be difficult. The important thing in this scale is wing
wear, and it increases with age, so record tears in the wings separate-
ly. A tear (or bird or lizard bite) can happen in a fresh specimen and
means little in this connection. Remember, you are using the scale to
estimate the relative age of the individual since eclosion, not to de-
scribe an exchange item! When you see a number of individuals of a
species on a particular day your notes might read, “all a” (suggesting
quite recent emergence), or “b-d, most c’ (suggesting about the
midpoint of the flight period or a little later), or “most d, one a”
(suggesting a second flight beginning as an earlier flight is ending),
and so on.

Words like “common” or “scarce” convey a poor idea of numbers;
mark-recapture techniques can give accurate population figures, but
the procedure is far too time-consuming for routine use. Some years
ago I devised a compromise system, more accurate and objective than
words, less tedious than population estimation; it is easily used, as
subsequent experience with it has shown. This abundance measure
is a logarithmic scale much like that of stellar magnitudes used in
astronomy, after which it was patterned. The scale records numbers
seen per hour, or the equivalent, as follows:

Scale Numbers

125-625 individuals seen per hour

25-125 e Oe ed
f= 25
5 z (sate

0.2- 1 per hour, or 1 seen per 1-5 hours

1 per 5-25 hours

1 per 25-125 hours

1 per 125-625 hours

1 per 625-3,125 hours

COnrn1mD wk WN Fe ©

The scale can be extended in either direction if needed (it rarely will
be): —1 would be the next commoner scale unit, and so on. Each
abundance unit represents ¥% the abundance of the preceding unit,
and a difference of one unit is about the minimum that can be accu-
rately perceived in routine observation.

228 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

In recording abundance take due note of highly localized species
(count only time spent in suitable habitats) and of flight periods (count
only time spent in appropriate months).

If you know your average rate of movement through an area, and
the average width of your “sweep —the distance you can effectively
survey on either side of your path (which will vary with the terrain
as well as your vision and knowledge of local butterflies)—then the
measure can be converted into a rough density measure of so many
per acre, or hectare, or whatever.

When only a single specimen is seen, its abundance should be
entered as “4 or rarer,’ “5 or rarer,’ etc., because catching one indi-
vidual cannot establish a time span.

The measure is crude, but about as accurate as possible in the cir-
cumstances. Remember, too, that any measure based on visual sight-
ing will generally underestimate true numbers.

Persistence pays. The whole idea is to continue observations for a
long time—several to many years—in all seasons. The following kinds
of data should be expected and sought:

Species list. The list will never be complete—something often not
realized—but with enough time you can approach completeness
about as closely as you wish. If you record the total accumulated
number of species you have found (S), and the total accumulated time
(in hours) you have spent observing (N), then the following relation
(Clench, 1968) will describe your results quite closely:

Se is the theoretical total number of species in your area, and K is an
adjustable constant, related to “collectability.”’

Powdermill Nature Reserve (of Carnegie Museum of Natural His-
tory) is an area of about 2,000 acres, located 9 miles south of Ligonier,
Westmoreland Co., Pennsylvania. Since its establishment in 1956 I
have worked on its butterflies as time allowed (little in recent years,
but intensively in the 1960’s). A total of 820 hours has been logged,
and 73 species recorded in that time: K = 59, and Se = 78 species.
In short, we have found about 94% of the species expected there,
with only 5 left to go (see Fig. 3). :

Calculating the “best fit” values of Se and K is difficult and com-
plicated, too much so for inclusion here. You can approximate them
fairly simply, however: draw a smooth, eye-fitted curve through your
graphed data points; pick two well spaced positions on this curve (or

VOLUME 33, NUMBER 4 229

80

Se = 78 species
1964 1968 Si
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1960 1962

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10

100 200

Total Man-Hours of Collecting (N)

Fic. 3. Collection curve of butterflies for Powdermill Nature Reserve (Westmore-
land Co., Pennsylvania). Spots represent cumulative man-hours of collecting (N) and
species recorded (S) at the close of each indicated year. The curve follows the formula:

oe 78( ew

two well spaced data points which touch it), substitute their S and N
values in the formula, and solve the two simultaneous equations for
Se and K. If your eye-fitted curve was well drawn, the results should
be fairly accurate.

A word of caution: the formula has ramifications and complications
that I have not mentioned and it should be used judiciously. By way
of example, Se actually represents the size of the “universe” being
sampled, and this may vary: a one-day universe is smaller than a one-
year universe, and that in turn is smaller than a two- or a ten-year
universe. A spring universe may be smaller than a midsummer uni-
verse and so on.

How many flight periods are there, and when? In any species the
timing, if not the number, of the flights varies from year to year, pre-

230 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

sumably because of variation in weather conditions. It takes several
years of observation to establish reliable average times.

One of the reasons for a diversity of flight periods among the species
of a community is to minimize competition for adult food—normally
flower nectar—and a thoughtful examination of your data in this light
could produce interesting results.

Population levels. Each species has an average level of abundance:
some are rare, others commoner, and some are abundant. In any given
area the frequency distribution of species according to abundance is
close to log-normal, that is, the frequency distribution approximates
a normal curve when abundance is plotted on a logarithmic scale.
Since the abundance scale (“0-8’’) described above is logarithmic, a
frequency distribution of the species based on their average abun-
dance according to that scale should approximate a normal curve. The
curve is truncated at the rare end, but the degree of truncation di-
minishes with prolonged study. Most species vary considerably in
their numbers from year to year, and again it takes a number of years
of observation to establish reliable average values. Some species may
have different abundance levels in different broods, and this varia-
tion, too, should be looked for.

Cycles of commonness and rarity are often present, but beware of
generalizing too much from them. Many cycles are transient, or not
truly cycles (having varying periods).

True regular cycles are uncommon. Eventually you should find in
some species a “boom and bust’ phenomenon. This is a season (or
flight) of exceptionally high numbers, followed by a crash, during
which you will see only a few individuals, or even none at all. A re-
turn to normal numbers soon follows. At Powdermill I have seen this in
a few species (Euphydryas phaeton, Polygonia comma, Epargyreus
clarus, Hesperia leonardus) over about 16 years of observation. In each
of these only one eruption was seen, ordinarily about 2 abundance
units above normal, followed by a crash of similar magnitude, with a
return to normal numbers the year after.

Habitat choices. With appropriate observation and recording, the
principal habitat and subordinate habitats of most of the species
should be learned in comparatively few years. Watch for species that
regularly occupy two or more different habitats, especially (a) for feed-
ing, and (b) for courtship, reproduction, larval growth, and inactive
(‘sleeping’) adult occupancy.

Territorialism. Many butterfly species are territorial. The subject
has scarcely been touched, and much remains to be learned. Persis-
tent observation should reveal territorial individuals if you are alert
to what the butterflies are doing. Absence of territorialism in a species

VOLUME 33, NUMBER 4 Dow

is harder to document, for individuals of territorial species are non-
territorial part of the time. Some species occupy territories only at
certain times of the day (such as Vanessa atalanta, in late afternoon).
In most territorial species males occupy the territories, and females
wander in search of them.

Life history data such as larval foodplants and adult food sources
should be noted. If you rear the early stages you can learn much more.

Long observation of an area will often show some transient species
that move into an area, live for one or a few seasons, and then die out.
At Powdermill, Nastra lherminieri arrived, lasted for several years,
then disappeared, and has not been seen since. Both Euptoieta clau-
dia and Hylephila phyleus established colonies that died out after a
single brood.

The Monarch, Danaus plexippus, is a special transient since it reg-
ularly moves into an area in the spring, raises a local brood or two,
then emigrates in the fall. The fall southward migration is conspicu-
ous and often spectacular. It is important to keep records of its size,
direction, and dates of start and finish. Just as important, however, is
the time of Monarch arrival in the spring, a far less documented event
because it is so inconspicuous. One can only note the date of first
spring sighting.

Inevitably, long-term observation will produce genuine resident
rarities, such as the celebrated Erora laeta, or strays from outside the
area—single individuals of non-resident species.

When you have studied an area intensively for several or many
years you are apt to find significant, non-random changes. They may
be changes in the trend levels of certain populations, but they could
be other things as well. As succession alters a meadow, grassland
species may change from commen to rare, or perhaps even disappear;
a species not seen before may move in and establish itself, perhaps
only temporarily, perhaps permanently.

LITERATURE CITED

CLENCH, H. K. 1949. Regional lists. Lepid. News 3: 15-16.

1968. A new butterfly for Powdermill. Powdermill Nature Reserve (Carnegie

Mus.), Educational Release no. 82: 3 p. [mimeographed].

1976. Nathalis iole (Pieridae) in the southeastern United States and the Ba-
hamas. J. Lepid. Soc. 30: 121-126.

FIELD, W. D., C. F. pos Passos & J. H. MASTERS. 1974. A bibliography of the catalogs,
lists, faunal and other papers on the butterflies of America north of Mexico arranged
by state and province (Lepidoptera: Rhopalocera). Smithsonian Contrib. Zool., no.
157: 104 p.

KIMBALL, C. P. 1965. The Lepidoptera of Florida. Gainesville (Fla. Dept. Agr., Div.
Plant Industry: Arthropods of Florida and Neighboring Land Areas, vol. 1): v +
363 p., ill.

Journal of the Lepidopterists’ Society
33(4), 1979, 232-238

DAILY FLIGHT PERIODS OF MALE CALLOSAMIA
PROMETHEA (SATURNIIDAE)!

MICHAEL E.. TOLIVER, JAMES G. STERNBURG, AND
GILBERT P. WALDBAUER

Department of Entomology, 320 Morrill Hall, University of Illinois,
Urbana, Illinois 61801

ABSTRACT. Flight periods of male Callosamia promethea were determined by
marking their positions in a large flight cage at hourly intervals from 0800 to 2000, and
recording the number in flight and the number moving from previous positions. Noc-
turnal flight activity was determined by marking male positions ca. 1 h after sunset and
again at sunrise. Flight activity occurs from 7 h before to 1 h after sunset and peaks 5-
2h before sunset. Preliminary observations of female pheromone release indicate that
pheromone release is synchronous with male flight activity, and peaks 4-1 h before
sunset. Pheromonal stimulation may be an important component in initiation of male
flight activity.

It has long been known that males of Callosamia promethea (Dru-
ry) are attracted to females by a pheromone during the afternoon,
usually between 1400 and sunset (e.g. Ferguson, 1972; Collins &
Weast, 1961; Eliot & Soule, 1902; Mayer, 1900). Rau & Rau (1929)
attempted to quantify the flight period of promethea by releasing both
bred and wild caught numbered males at various distances from caged
females and noting the time of arrival of males. They found a peak of
activity from 1600-1640 Central Standard Time at St. Louis, Missouri,
with 14 of 33 recaptured males returning during this period. They
also observed one male arriving at dawn with males of Hyalophora
cecropia (L.), though they did not state to which species (female ce-
cropia or female promethea) the promethea male was attracted. In
all of the above work, male flight periods were apparently determined
by watching caged females and observing time of male arrivals. Thus,
these observations do not establish conclusively that flight activity of
male promethea is limited to the afternoon. This aspect of behavior
can be determined only be watching males throughout the day.

Because of our studies on mimicry, involving the release and re-
capture of variously painted promethea males (Waldbauer & Stern-
burg, 1973; Sternburg et al., 1977), it became necessary to define more
precisely the flight activity of males. If male promethea were noctur-
nal as well as diurnal, nocturnal predators could account for the dif-
ferential recapture of mimetic vs. non-mimetic color patterns ob-
served by Sternburg et al. (1977). The non-mimetic yellow pattern

' Part of the Ph.D. dissertation presented by the first author to the Department of Entomology, University of
Illinois.

VOLUME 33, NUMBER 4 233

may be more conspicuous at night than the black mimetic pattern. If
males fly only in the afternoon, then exposure to diurnal predators
will be less than if males were to fly in both the morning and after-
noon.

MATERIALS AND METHODS

We used freshly emerged male and female promethea reared on
wild black cherry (Prunus serotina Ehrh.) or tuliptree (Liriodendron
tulipifera L.) at Urbana, Illinois. Adults emerged from pupae allowed
to overwinter in an outdoor insectary. These stocks originated from
wild populations from the vicinity of Charleston, Illinois or Medary-
ville, Indiana.

Males were individually distinguished by white numbers painted
on the ventral surface of the hindwings. They were released into a
flight cage (2.36 m x 1.83 m X 2.36 m) containing a wild black cherry
tree (ca. 1.8 m tall) and a recording thermograph. They were observed,
their positions marked, and the number in flight (if any) noted at
hourly intervals from 0800 to 2000 (all times Central Daylight time)
each day from 8 to 11 July 1977. There were 16 males in the cage on
8, 10, 11 July, and 15 males on 9 July. New males were added to the
cage at 1100 on 10 and 11 July. They were allowed to settle; then
their positions were marked. At the next hourly observation, their
positions were noted along with the positions of males which had
been present all day. Only one of the 15 males added to the cage this
way moved during that hour. This four-day intensive observation pe-
riod was supplemented by less regular observations from 3 June to 8
July that are not included in the table.

Night activity in 1977 was determined by mapping the positions of
males at sunset and again at sunrise. When it was found that no move-
ment took place between sunrise and 0900, the morning observations
were changed to between 0700 and 0900. These observations were
made on eight nights in June, 1977, using a total of 115 males. Because
a higher percentage of these males moved at some time during the
night than expected (22.8%), nighttime observations were repeated
in 1978.

The flight cage was moved to a new location over a small, dense,
low-growing barberry (Berberis) cultivar in an effort to provide males
with sheltered resting sites. However, most males continued to rest
on the sides and top of the cage (see below) rather than in vegetation.
Male positions were marked between 2100 and 2200 and again be-
tween 0500 and 0600 the following day. Sixty-two observations on 30
males were made between 29-31 May and between 5-7 June 1978.

234 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

RESULTS AND DISCUSSION

Table 1 shows that most diurnal flight activity of male promethea
occurs during the afternoon between 1300 and 2000. The peak of
activity occurs from 1600 to 1700, 4 to 3 h before sunset. There is
some activity in the morning, but we believe this may be an artifact
of confinement in the flight cage. The natural resting site of wild
males is presumed to be in vegetation, but they often rested on the
sides or top of our cage. For example, on the morning of 8 July, 13 of
16 males were resting on the sides or top of the cage and not in the
tree. Seven of these 13 males moved during the period 1000-1100
(70% of all males moving during these hours on all four days). Of
those moths moving, three moved 0.3 m or less. Of the four remaining
males, three moved from exposed positions on the top of the cage to
sheltered positions on the cage sides, while the fourth male moved
from an exposed position on one side to a sheltered position on
another side. We believe that males which moved in the morning
were shifting from positions exposed to the sun, possibly to avoid
increasing temperatures. On 8 July, when most of these males moved,
the temperature reached 31.1°C at 1400, the highest recorded for the
four-day period, and was already 29.4°C at 1100. During the period
1000-1100, 8 July, the temperature rose 2.8°C, the largest increase in
temperature in one hour during the period 0800-1100 on any of the
four days. The maximum temperature for the other three days during
the period 0800-1100 was 27.2°C, while the largest hourly increase
in temperature was 2.2°C. Other morning movements were similar in
that males moved from exposed to sheltered positions, although not
all males in exposed positions moved. None of the males which rested
in the tree (five males over four days) moved before 1400. Only three
males were observed to fly in the morning, one because it had been
disturbed (Table 1). Males were observed in flight primarily at 1600
and 1700, corresponding well with the number of males observed to
change their positions.

In 1977, 23 of the 115 males observed to determine nocturnal move-
ments were eliminated from the data: 11 because they could not be
found at sunset to mark their positions, one because it flew to an
unknown location when it was disturbed as its position was marked,
and 11 because they died during the night. Of the remaining 92 males,
21 (22.8%) moved during the night. This is a higher percentage of
movement than expected. The 1978 observations indicate that this
high percentage was the result of marking male positions too early in
the evening. In 1977, male positions were marked at sunset, which
occurs shortly after 2000 in Urbana at the time of year these obser-

235

VOLUME 33, NUMBER 4

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236 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

vations were made. Table 1 shows that nearly 20% of males moved
between 1900 and 2000. In 1978, male positions were marked be-
tween 2100 and 2200, approximately one hour after sunset. When this
was done, only two males (3.2%) were observed to move during the
night. Both of these males exhibited wing fluttering behavior when
their night positions were first marked, and did not fold their wings
and rest quietly during the 20 to 30 minute period required to mark
all male positions. Thus it seems likely that these two males moved
shortly after we left the flight cage for the night.

Preliminary observations of virgin female promethea indicate that
peak flight activity of males corresponds roughly to peak pheromone
release by females. Sixty-eight females were watched in the early
afternoons of 16, 29, 30 June and 1, 5, and 6 July for extrusion of an
abdominal gland, which, on the basis of male behavior, is obviously
the source of the sex pheromone. Females began extruding this gland
between 1400 and 1500 (three females) and by 1700, 59 of the 68
females had extruded it. Of the remaining nine females, eight ex-
tended the gland during the next hour; while one had not extruded
her gland by 1800 when observations ceased. Observations of some
of these and of other females during the morning and early afternoon
(before 1400) indicated that the abdominal gland is never extended
between 0800 and 1300. Thirty-three females were watched on the
evenings of 14, 15, 16 and 22 June to observe times of retraction of
the abdominal gland. One female retracted the gland between 1800
and 1900, 12 retracted it between 1900 and 2000, and 20 retracted it
between 2000 and 2100. At this time of year sunset occurs at Urbana
between 2023 and 2026. Thus, female promethea stop releasing pher-
omone before twilight ends. To make sure nocturnal release did not
occur, 12 females were watched all night on 22 June. Observations
were made hourly up to 2200, and every 2 hours thereafter until 0600,
23 June. Females were observed with a flashlight covered by a red
cloth to minimize disturbance. At no time did any of them extrude
their abdominal glands.

Collins & Weast (1961) noted that certain atmospheric conditions
could cause males to fly as early as 1300. Their times are presumably
standard times, while ours are daylight times; thus our 1400 corre-
sponds to their 1300. We noted that females tended to extrude their
glands earlier on cloudy days, but did not make enough observations
under appropriate conditions to quantify this.

Skinner (1914) found that male Callosamia angulifera (Walker)
were attracted to female C. promethea between 2000 and 2100. In
the course of our release-recapture experiments with painted male
promethea in Urbana, we attracted a male C. angulifera with a female

VOLUME 33, NUMBER 4 a3 Th

promethea sometime between 1930 8 August and 1900 9 August,
1977, and another male angulifera sometime between 1300 12 June
and 1900 13 June, 1978. Ferguson (1972) notes that female C. angu-
lifera attract males between dusk and midnight, with a peak activity
at 2200. The slight overlap of promethea pheromone release and an-
gulifera flight activity may account for this interspecific attraction.
The three species of Callosamia appear to be reproductively isolated
by their differing pheromone release and male flight periods (Fer-
guson 1972) but the ability to attract males of one species with females
of another indicates that this isolating mechanism occasionally breaks
down.

Brown (1972) suggested that males of Callosamia securifera (Maas-
sen) did not need a pheromone stimulus to start flight activity, which
occurs from 1000 until 1500 in this species. Our observations indicate
that pheromonal stimulation may be a component of flight initiation
in promethea: For example, on 9 June at 1425, 10 males were in the
flight cage in the same positions they had occupied since 0940 that
morning. Two females were in a cage upwind close by and neither
had been observed to extrude the abdominal glands all day. Between
1425 and 1435 most of the males began quivering, a characteristic
behavior exhibited immediately before afternoon flights, and four
males actually flew. We then checked the female cage and found that
one female was extruding her abdominal gland. The second female
extruded her gland 10 min. later. By 1500, all but 2 of the males had
flown. By moving female cages downwind, we could often cause
males in the flight cage to settle. No experiments were tried where
females were absent to see if males would fly without pheromonal
stimulation, as the area in which the flight cage was located possesses
a wild population of promethea and the possibility of stimulation by
a wild female could not be excluded.

Other work by Jeffords, Sternburg and Waldbauer (in prep.) indi-
cates that male promethea do fly in the afternoon without pheromonal
stimulation. They released painted males of promethea at Allerton
Park, Piatt Co., Illinois, where either no or a very small wild popu-
lation of promethea exists, and then recaptured these males one day
later in traps baited with virgin females. Wing damage analysis and
the ratio of mimetic to non-mimetic painted male recaptures indicated
that these males had flown the previous day when no females were
present. However, the possibility of stimulation by wild females can-
not be entirely discounted.

CONCLUSIONS

Male promethea are diurnal fliers, with a peak activity between

238 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

1500 and 1800, or between 5 and 2 h before sunset. The slight amount
of morning movement is believed to be an artifact of confinement in
a flight cage. Nocturnal movement does occur, but it is limited to a
period less than 2 h after sunset and is thus more appropriately con-
sidered crepuscular. Thus males are exposed in flight to predators
mainly in the afternoon.

Female sex pheromone release is apparently confined to the after-
noon between 1400 and 2100 and peaks between 1600 and 1900 (4 to
1 h before sunset). No nocturnal or morning release, as evidenced by
extrusion of the abdominal gland, was noted. Male flight activity may
be initiated by reception of the female pheromone, and/or it may be
the result of endogenous rhythms or other environmental stimuli.

ACKNOWLEDGMENTS

We thank Dr. C. T. Maier for the use of the flight cage, and Dr. W.
H. Luckmann and the Illinois Natural History Survey for the use of

the experimental area and facilities for rearing promethea. This work
was supported by NSF Grant DEB 76-18618 to Waldbauer and Stern-
burg.

LITERATURE CITED

BROWN, L. N. 1972. Mating behavior and life habits of the sweet-bay silk moth (Cal-
losamia carolina). Science 176: 73-75.

Couuins, M. M. & R. D. WEAST. 1961. Wild silk moths of the United States. Collins
Radio Co., Cedar Rapids, iii + 138 p.

EvIoT, I. M. & C. G. SOULE. 1902. Caterpillars and their moths. Century Co., N.Y.
302 p.

FERGUSON, D. C. 1972. Bombycoidea, Saturniidae (in part). In R. B. Dominick, et al.,
The moths of America north of Mexico, fasc. 20.2B: xxi + 155-275, pls. 12-22.

MAYER, A. G. 1900. On the mating instinct in moths. Psyche 9: 15-20.
RAvu, P. & N. Rau. 1929. The sex attraction and rhythmic periodicity in the giant
saturniid moths. Trans. Acad. Sci. St. Louis 26: 81-221.
SKINNER, H. 1914. Callosamia promethea and angulifera (Lepid.). Entomol. News
25: 468-469.

STERNBURG, J. G., G. P. WALDBAUER, & M. R. JEFFORDS. 1977. Batesian mimicry:
selective advantage of color pattern. Science 195: 681-683.

WALDBAUER, G. P. & J. G. STERNBURG. 1975. Saturniid moths as mimics: an alter-
native interpretation of attempts to demonstrate mimetic advantage in nature. Evo-
lution 29: 650-658.

Journal of the Lepidopterists’ Society
33(4), 1979, 239-244

COMPOSITES AS HOST PLANTS AND CRYPTS FOR
SYNCHLORA AERATA (GEOMETRIDAE)

MIKLOS TREIBER!

Department of Botany, University of North Carolina,
Chapel Hill, North Carolina 27514

ABSTRACT. Thirteen genera of Asteraceae have been recorded as host plants of
the geometer moth Synchlora aerata. These flower-feeding larvae not only live on
their food plants but also use the host plants as sources of raw material to disguise
themselves. The larvae cut off flower parts, entire flowers, and even entire inflores-
cences and attach them to spiculiferous processes on their dorsal surface. If this floral
covering is removed the larvae immediately replace the camouflage; furthermore, the
larvae maintain the effectiveness of the covering by replacing withered floral parts with
fresh ones. Adaptation to the use of host plant material as camouflage enables the larvae
to exploit, with greater safety from predators a greater range of potential food plants.

Little is known of the life history of the larvae of geometer moths.
On the basis of their structure and behavior in relation to concealment
from predators, the larvae so far described can be divided into three
distinct groups: 1) those that are slender and twig-like; 2) those with
moderate dorsolateral processes bearing specialized hooks for the at-
tachment of plant fragments as an aid to concealment, and 3) those
with large dorsolateral processes not specialized for the attachment
of plant matter (Ferguson, 1969). Of interest here are the behavioral
characteristics and host plant relations of Synchlora aerata with the
Asteraceae (=Compositae). Synchlora aerata belongs to the second
of the above groups.

Larvae of S. aerata were observed on, and collected from, two
species of Liatris on the coastal plain of North Carolina in Carteret
Co., and on three species of Solidago on the piedmont of North Car-
olina in Orange Co. They have also been reported on seven other
composite genera (Table 1). Larvae of S. liquoraria liquoraria and S.
frondaria have also been reported on composites (Table 1).

METHODS AND MATERIALS

The larvae were collected and transplanted to terraria where their
feeding behavior, and their use of plant fragments as camouflage,
could be observed. The response of the larvae was studied under the
following conditions: 1) when camouflage was removed by artificial
means, and 2) when host plants were changed. In the latter case,

1 Present address: U.S. Army Engineer Topographic Laboratories, Research Institute, Center for Remote Sensing,
Fort Belvoir, VA 22060.

240 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TABLE 1. Summary of the species of Synchlora and their known composite host
plants.

Species Host plants Citation
S. aerata (Fabricius) Ambrosia sp. Ferguson (1969)
Dyar Coreopsis sp. Ferguson (1969)
Erigeron canadensis Ferguson (1969)
Rudbeckia hirta Ferguson (1969)
Rudbeckia sp. Ferguson (1969)
Ageratum sp. Anonymous (1890)
Eupatorium sp. Anonymous-(1890)
Liatris graminifolia Treiber*
L. spicata var. resinosa Treiber*
Solidago canadensis Treiber*
S. nemoralis Treiber*
S. pinetorum Treiber*
Aster sp. Dyar (1960)
S. liquoraria liquoraria Artemisia californica Comstock & Dammers
Guenee (1937)
Solidago sp. Comstock & Dammers
(1937)
S. frondaria Guenee Bidens sp. Ferguson (1969)
Pluchea odorata Ferguson (1969)
Chrysanthemum sp. Kimball (1965)

* First report.

larvae feeding on Liatris were transplanted to Solidago and vice ver-
sa. One individual was successfully reared and was the basis for iden-
tification as Synchlora aerata. Voucher specimens of both the larval
stage and adult stage have been deposited into the U.S. National Mu-
seum, Dept. of Entomology, Smithsonian Institution.

FEEDING AND LOADING BEHAVIOR

Observations on feeding and camouflage-loading behavior were
first made on Liatris spicata var. resinosa (Fig. 1). In each case, the
larvae began dissecting individual flowers by cutting the corolla tube
vertically between lobes for a distance of 1-2 mm, then cutting per-
pendicularly to the long axis of the corolla tube for nearly its entire
circumference. The corolla was further dissected into fragments with
1 to 3 lobes. The cut edge of each section was passed through the
larva’s mouth, and carefully attached to a spiculiferous process on its
back (Fig. 2). As the fragments were passed through the mouth of the
larva, a mucilaginous substance was secreted by the larva onto the
fragments, this substance seems to play a role in maintaining turgor
in these fragments. Next, the style (and in a few cases pappus hairs)
were bitten off and also treated as described above. The larva then

VOLUME 33, NUMBER 4 241

ye PLU DOR CC ko Ce CO
1 2 3

Fic. 1. Macrophotographs of the larva of Synchlora aerata on Liatris spicata var.
resinosa; (A) inflorescence stalk of L. spicata var. resinosa with arrow pointing to the
location of a fully camouflaged larva and (B) a comparison of the size of an inflorescence
and of a flower of L. spicata var. resinosa with a fully camouflaged larva.

devoured the anther sacs and pollen, which apparently provide its
chief source of nourishment, although some corolla and stylar tissue
was also observed being ingested. In no case was the ovary of the
flower damaged.

Even when the disguise is complete, maintenance is required. Two
kinds of maintenance are recognized: 1) the addition of a mucilagi-
nous substance, which appears to maintain turgor in the plant frag-
ments, and 2) the replacement of “older” missing, lost, or wilted tis-
sue with “new tissue. Apparently, the “old” tissue is either ingested
by the larvae or discarded at some point and subsequently replaced.
Thus the loading process is more or less continuous.

The feeding behavior on Solidago flowers was similar to the
behavior observed when feeding on Liatris, that is, the principal
nourishment was derived from the anthers and pollen, as well as from
occasional feeding on stylar and corolla tissue. However, the camou-
flage-loading behavior was different. On Solidago the floral dissecting
process described above was omitted. Instead of dissecting corolla
fragments, the larva bit off entire inflorescences and attached them to
its back in the manner described for Liatris (Fig. 2). Apparently, frag-

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

242

VOLUME 33, NUMBER 4 243

ments from the larger flowers of Liatris were sufficient to provide a
camouflage; however, in the small-flowered Solidago species, flower
fragments were insufficient to effect a viable camouflage. Both types
of loading behavior, described above, were exhibited by the same
larva.

In order to better understand the loading behavicr and the rela-
tionship between feeding and loading, three larvae were artificially
denuded. The larvae immediately began to feed and load alternative-
ly until their camouflage was complete. In similar experiments on
Chrysopa slossonae (Neuroptera) Eisner et al. (1978) found that the
relative priority the larvae give to these two activities is variable and
is dependent on the degree of satiation. However, in the case where
the larvae were denuded and hungry, the larvae divided their time
between loading and eating. In addition, denuded larvae, as well as
larvae with a complete disguise, were transplanted from Liatris to
Solidago and vice versa. The denuded larvae again began to feed and
load until the disguise was complete. The larvae, with a complete
camouflage of plant material from the first host plant, were observed
to replace these plant fragments with floral material from the second
host plant. Replacement proceeded in the same manner as observed
in the maintenance process. Whether or not the difference in host
plant was recognized by the larvae can only be conjectured; however,
the result was a complete change of the larvae’s covering to match
the new host plant.

DISCUSSION

The adaptation of loading plant fragments, flowers, and/or inflores-
cences as a disguise by the larvae of Synchlora aerata effects an
apparently successful camouflage against winged predators. More im-
portantly, this adaptation enables the larvae to feed on a variety of
related host plant species rather than being restricted to a specific
host plant, as in the case with adaptations to special marking or col-
oration, due to predator pressure. This adaptation is viewed as an
important evolutionary adaptation enabling the larvae of S. aerata to
exploit a greater range of potential food plants. This conclusion is
supported by the relatively large number of composite genera that
have been recorded as host plants.

oo

Fic. 2. Macrophotographs of the larva of the geometer moth Synchlora aerata on
the goldenrod Solidago canadensis; (A) showing the orientation and arrangement of
goldenrod inflorescences attached to the dorsal surface of a larva; (B) dorsal view of
larva after removal of a portion of its camouflage, and (C) lateral view of larva after
removal of a portion of its camouflage.

244 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

ACKNOWLEDGMENTS

I thank Drs. C. R. Bell, Guy L. Nesom, and J. N. Rinker for their
critical review of this manuscript and Drs. John C. Downey, Herbert
Neunzig, and Thomas Eisner for their enlightening discussions and
encouragement. Linda S. Treiber and Mary L. May have my appre-
ciation for their field assistance; and I thank Paul Marx for his pho-
tographic advice and assistance.

LITERATURE CITED

ANONYMOUS. 1890. Some of the bred parasitic Hymenoptera in the national collection.
Insect Life 2: 348-353.

COMSTOCK, J. A. & C. M. DAMMERS. 1937. Notes on the early stages of three California
moths. So. Calif. Acad. Sci. Bull. 36: 68-78.

Dyar, H. G. 1900. Life histories of North American Geometridae XIII. Psyche 9:
93-94.

EISNER, T., K. Hicks & M. EISNER. 1978. “Wolf-in-sheep’s clothing” strategy of a
predaceous insect larva. Science 199: 790-794.

FERGUSON, D. C. 1969. A revision of the moths of the subfamily Geometridae of
America north of Mexico (Insecta, Lepidoptera). Peabody Museum of Natural His-
tory Bull. 29. 1-215 p.

KIMBALL, C. P. 1965. Lepidoptera of Florida. Arthropods of Florida and neighboring
land areas, Vol. 1: 363 p., illus. Div. of Plant Industry, Florida Dept. Agric., Gaines-
ville.

Journal of the Lepidopterisis’ Society
33(4), 1979, 244

A NEW METHOD OF INDUCING COPULATION IN PHYCIODES THAROS
(NYMPHALIDAE)

While engaged in breeding experiments using various populations of Phyciodes thar-
os Drury, I happened on a method of inducing copulation that may be widely appli-
cable to other butterfly species. It proved extremely difficult to achieve matings by the
hand-pairing technique or in small cages using several population cultures of P. tharos
in my laboratory. The difficulty appeared to be both an unusually low level of courtship
activity in the males and an unusually low proclivity toward acceptance by the females.
However, I noticed that stray males that had escaped from the mating cages and flown
to a large screened window often showed greatly increased aggressive behavior and
sexual approaches toward each other. Females placed on the screen near courting
males still refused to mate, but when they were restrained by holding the wings to-
gether over the back with a pair of flat forceps, the males were often able to copulate.
Greater success was achieved by stroking the female’s abdomen on the male’s antennae
to elicit repeated copulation attempts and by moving the female’s abdomen to bring
her genitalia into contact with the male’s. If the female was released at this point, she
still attempted to avoid copulation and would often dislodge the male by her struggles.
I had better success by pinching the forceps handle with a spring-type clothespin and
putting the clothespin across the mouth of a small jam jar. The quiescent male then
hung from the female’s abdomen until copulation was complete, when he dropped to
the bottom of the jar. This method may prove to give better results than either cage or
hand-pairing for a number of difficult species.

CHARLES G. OLIVER, R.D. 1, Box 78, Scottdale, Pennsylvania 15683.

Journal of the Lepidopterists’ Society
33(4), 1979, 245-247

A NEW SUBSPECIES OF ORGYIA LEUCOSTIGMA
(LYMANTRIIDAE) FROM SABLE ISLAND, NOVA SCOTIA

KENNETH NEIL
Dept. of Biology, Dalhousie University, Halifax, Nova Scotia, Canada B3H 4J1

ABSTRACT. A new subspecies of Orgyia leucostigma (J. E. Smith) from Sable
Island, Nova Scotia is figured and described.

Sable Island is a small, sandy, crescent-shaped island located 150
mi due east of Halifax, Nova Scotia. Until 1975, the Lepidoptera fauna
was poorly known with only 23 species (Noctuoidea) listed as occur-
ring on the island (Howden, 1970). Extensive collecting over the last
three years by Barry Wright of the Nova Scotia Museum increased
this number to 58 (Neil, 1977), adding much to the local knowledge
of the Lepidoptera of the island.

One of the more interesting captures taken during the course of this
study was a small series of seemingly aberrant male Orgyia leuco-
stigma (J. E. Smith) collected in 1976. These specimens had most of
the brownish-black ground color of O. leucostigma plagiata (Walker)
replaced by a dull rusty brown and were very indistinctly marked.
Since only a few specimens were available for study, no immediate
work was done on them, and a concentrated effort to obtain wild
specimens and egg masses for breeding purposes was made in 1977.
Numerous specimens subsequently reared under laboratory condi-
tions were similar to the wild specimens collected on the island. It
then became evident that this form is genetically different and rep-
resents an undescribed subspecies endemic to Sable Island, as men-
tioned by Ferguson (1978), who also illustrated a male of this sub-
species (1978: 85, pl. A, Fig. 13).

Orgyia leucostigma plagiata is known with certainty from only
Nova Scotia, Prince Edward Island, and New Brunswick, and repre-
sents a Pleistocene relict endemic to the Gulf of St. Lawrence region
(Ferguson, 1978). The larvae are extremely general feeders, having
been recorded from almost every kind of tree and shrub, and are
regarded as a serious pest in Nova Scotia, especially on balsam fir
trees.

Orgyia leucostigma sablensis Neil, new subspecies

Description. Male: Upperside of forewing dull rusty brown. Markings and lines
as in plagiata but generally much more diffuse and indistinct. Median space rusty
brown with varying amounts of blackish scales from anterior apex of discal spot to
above second anal vein. Light grey of median area found in plagiata generally absent,
or if present, reduced to a small patch at costa. Basal third of forewing also rusty brown
with the darker brown shades of plagiata reduced to a small patch at costa. Discal dot

246 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fics. 1-4. Orgyia spp. 1, O. leucostigma sablensis 3, holotype; 2, O. leucostigma
sablensis 9°, allotype; 3, O. leucostigma plagiata 6, Lunenburg, Nova Scotia, 21 Aug.
1976, B. Wright; 4, O. leucostigma plagiata 2°, McNabs Island, Halifax Co., Nova
Scotia, 5 Sept. 1905, J. Perrin.

obscure and indistinct. Antemedial line very diffuse, lighter brown in color than sur-
rounding basal area. Outer third of forewing also rusty brown with a variable amount
of blackish scaling present distad of postmedial line. Dark color narrow at apex wid-
ening substantially at tornus. Dark brown spot immediately distad of postmedial line
reduced or obsolete. Medial and postmedial lines present but diffuse. Fringe concol-
orous.

Upperside of hindwing solid rusty brown with fringe concolorous or slightly lighter.

Underside of both wings rusty brown, somewhat lighter along the posterior margin
of the forewing. A darker brown postmedial line and discal dot present on both fore-
and hindwings of some specimens.

Vestiture with coloration as in ground of wings. No visible differences from plagiata
in antennae, palpi, or vestiture of head and body.

Length of forewing: holotype male, 14 mm; of male paratypes, 10-15.5 mm.

Female. Small wing pads present. Coloration of body pale grey, almost white both
dorsally and laterally. Antennae, palpi, and other external structures as in plagiata.
Vestiture also as in plagiata.

Male and female genitalia. Identical to those of plagiata.

Types. Holotype: 3, Meteorological Station, Sable Island, Nova Scotia, 11 Sept.
1977, B. Wright (Fig. 1). Allotype: 2, Main Station, Sable Island, Nova Scotia, 26 July
1976, B. Wright (Fig. 2), reared ex larvae on Bayberry (Myrica pensylvanica Loisel),
emerged 10 Aug. 1977. Paratypes: 2 6, West Light, Sable Island, Nova Scotia, 3 Aug.

VOLUME 33, NUMBER 4 247

1976, B. Wright; ¢, same data as above, but taken 6 Aug. 1976; 3 3, same data as
allotype, emerged 10 and 13 Aug. 1976; 17 5, 98 2, same data as holotype, but collected
on 10 Sept. 1977, reared ex ova on artificial medium (Shorey and Hale, 1965), emerged
6-20 Mar. 1978. Holotype and allotype have been deposited in the Canadian National
Collection. Paratypes have been deposited in the collections of the U.S. National Mu-
seum, Nova Scotia Museum, and the K. Neil collection.

Distribution. The type locality represents the entire known distribution of sablen-
sis.

Early stages. Larvae structurally identical to those of plagiata found on mainland
Nova Scotia, but with a tendency to differ in color. A large proportion of the larvae, as
many as 35-40%, resemble those of Orgyia leucostigma intermedia Fitch of the eastern
United States, in having red heads, a grey dorsolateral band, blackish areas of the body
much reduced, and whitish tufts. This type of larva is quite rare on mainland Nova
Scotia and comprises only about 1% of the population (Ferguson, ibid.).

Foodplants. A general feeder, like other subspecies of leucostigma. Larvae col-
lected from blueberry and cranberry (Vaccinium sp.), iris (Iris versicolor L.), bayberry
(Myrica pensylvanica Loisel), rose (Rosa sp.), and several species of grasses and
sedges.

Remarks. Sablensis is the first endemic moth to be recorded from Sable Island.
This subspecies also represents a Pleistocene relict that survived glaciation on Sable
Island and has since evolved into this unique insular subspecies. The flight period
corresponds to that of mainland plagiata with adults occurring from late July to mid-
September.

ACKNOWLEDGMENTS

I thank Barry Wright of the Nova Scotia Museum for his numerous
comments and suggestions made during the preparation of this paper
and for his review of the final manuscript, and D. C. Ferguson of the
Systematic Entomology Laboratory, U.S. Dept. of Agriculture, Wash-
ington, D.C. for his review of the final manuscript, and Mary Primrose
of Dalhousie University for photographing the types.

LITERATURE CITED

FERGUSON, D. C. 1955. The Lepidoptera of Nova Scotia: (Macrolepidoptera): Nova
Scotia Mus. Sci. Bull. 2: 214 p.

1978. In Dominick, R. B. et al. 1978. The moths of America north of Mexico,
Fasc. 22.2, Noctuoidea (in part): Lymantriidae.

HOwWDEN, H. F., MARTIN, J. E. H., BOUSFIELD, E. L., & D. E. MCALLISTER. 1970.
Fauna of Sable Island and its zoogeographical affinities—a compendium. Nat. Mus.
Natl. Sci. Publ. Zool. No. 4: 1-45.

NEIL, K. A. 1977. An Annotated List of the Macrolepidoptera of Sable Island. Proc.
N.S. Inst. Sci. 28: 41-46.

SHoREY, H. H. & R. L. HALE. 1965. Mass-rearing of the larvae of nine noctuid species
on a single artificial medium. J. Econ. Ent. 58: 522-524.

Journal of the Lepidopterists’ Society
33(4), 1979, 248-253

CALLOPHRYS NIPHON (LYCAENIDAE) IN ALBERTA
WITH NOTES ON THE IDENTIFICATION OF
C. NIPHON AND C. ERYPHON

JAMES D. REIST!
Department of Zoology, University of Alberta, Edmonton, Alberta, Canada, T6G 2E9

ABSTRACT. The known range of Callophrys niphon clarki is extended 185 miles
west into the province of Alberta, Canada with the capture of several specimens in
May 1977 and May 1978.

Atypical specimens of Callophrys eryphon eryphon, a species closely related to C.
niphon, possess a mid-cell and an end of discal cell bar thus invalidating an important
character heavily relied upon for identifying these butterflies. Some atypical specimens
of C. niphon have a smeared or rarely absent mid-cell bar. Alberta C. niphon show
great variation in other characters including those used for identification purposes in
keys of major works. Thus, extreme caution is warranted when identifying these two
species using existing taxonomic and descriptive literature.

On the basis of identification of two specimens confirmed by Dr.
C. D. Bird and three additional specimens (of poorer quality) con-
firmed by Mr. J. D. Lafontaine, Callophrys (Incisalia) niphon clarki
(Freeman) is herein reported to occur in jackpine woods in central
Alberta (54°5'N, 113°50’W) (Figs. 1 and 2).

Hooper (1973) reports the northwestern range of C. niphon clarki
to be “the Loon Lake area,’ Saskatchewan (about 35 miles east of the
Alberta-Saskatchewan border at 54°N). Other references to the west-
ern range list it simply as southern Manitoba (Brown, 1957; Clench,
1961; Howe, 1975).

The initial collection consists of two specimens in good condition
taken along a cutline in a jackpine wood 7% mi east and 2% mi south
of Clyde, Alberta on 7 May 1977. The immediate topography consists
of low rolling hills with a shallow dip to the west. The elevation of
the area is approximately 2,100 ft. Vegetation is almost totally jackpine
(Pinus banksiana) with sparse, low shrub and herbaceous under-
growth. Aspen poplar (Populus tremuloides) and shrubs such as sas-
katoon (Amelanchier alnifolia) and rose (Rosa sp.) have regenerated
along road allowances, cutlines and occasional clearings. Callophrys
augustinus augustinus (Westwood) and Callophrys polios obscurus
Ferris and Fisher were also taken at this time.

On 4 June 1977 five additional specimens of C. niphon, in poor
condition were collected at West Bridges Lake, 2% mi north and 6%
mi east of Clyde. This site is approximately 5 mi NNW of the first
collection area and in the same type of topography and vegetation

' Current address: Department of Ichthyology and Herpetology, Royal Ontario Museum, 100 Queen’s Park, To-
ronto, Ontario, Canada, M5S 2C6.

VOLUME 33, NUMBER 4 249

community. These specimens were all taken on a moist, sandy beach
on the west side of the lake. The immediate vegetation was an aspen
poplar-shrub wood bordering the lake for about 200 yards before giv-
ing way to a jackpine community such as described above. C. polios
was also taken here.

One additional worn specimen was taken by Mr. T. W. Thormin on
6 June 1977 six mi east of Redwater, Alberta; that is, 22 mi SE of the
first collection site.

On 6 May 1978 three specimens of C. niphon were taken at the
first site. On 7 and 13 May 1978 many additional (about 40) specimens
were taken in the jackpine wood on the west side of West Bridges
Lake. Both sexes were in flight but oviposition was not observed. C.
polios and C. augustinus were flying at both sites as well. T. W.
Thormin took several C. niphon near Bonnyville, Alberta on 28 May
1978.

Fig. 1 shows the known localities, to date, of C. niphon clarki in
Alberta. The widespread occurrence of C. niphon, both in space and
time, suggests this species is well established in the province prob-
ably as a breeding population(s). Jackpine, preferred foodplant of C.
niphon, is found over large areas of central east and northeastern
Alberta (Fig. 1); thus additional collecting in these areas may greatly
extend our knowledge of the provincial range of this species. The
collections herein reported extend the known range of C. niphon
clarki approximately 185 mi further west than previously determined
(Hooper, 1973).

Fig. 1 also shows the Alberta distribution of Callophrys (Incisalia)
eryphon eryphon (Boisduval), a closely related congener of C. ni-
phon. The preferred foodplant of C. eryphon (Pinus contorta—lodge-
pole pine) occurs primarily in montane and submontane areas of west-
ern Alberta. However, lodgepole pine and jackpine distributions
overlap considerably in central Alberta. This raises the possibility of
sympatry of these two butterflies in Alberta. With sympatry comes the
possibility of confusion in identification and the potential for hybrid-
ization. The remainder of this paper will address the first of these
possibilities.

IDENTIFICATION OF C. eryphon AND C. niphon

Identification of the initial C. niphon specimens proved to be dif-
ficult using the available literature. Examination of six specimens of
both C. niphon and C. eryphon from the Biosystematics Research
Institute, the C. eryphon collections of both the University of Alberta
and University of Calgary, and a consideration of the 1978 Alberta C.
niphon showed that the characters commonly used for identification

250 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

°
i—]
N
=

Fics. 1, 2. 1, Outline map of Alberta, Canada showing known localities of
Callophrys eryphon eryphon (triangles) (C. D. Bird, pers. comm.) and C. niphon clarki
(closed circles). Also shown are the eastern limits of Pinus contorta latifolia (dotted

line) and western and southern limits of Pinus banksiana (toothed line) (both after

VOLUME 33, NUMBER 4 Zeal

are not constant in specimens from different areas. Variation com-
monly is towards the other species, leading Brown (1957) to question
the validity of two species.

Most authors use three characters for identification: presence of
both a mid-cell and end of cell dark bar transversely crossing the
discal cell on the underside of the forewing in C. niphon (a and b in
Figs. 2 and 4), while typical C. eryphon lack the mid-cell bar (Fig.
3), (Clench, 1961; Hooper, 1973; Howe, 1975). The submarginal dark
chevrons on the underside of the hindwing in C. niphon are shallowly
angled compared to the same on C. eryphon (c in Figs. 2 and 3),
(Brown, 1957; Clench, 1961; Hooper, 1973; Howe, 1975). C. niphon
also has more gray scaling on the submarginal line of the underside
hindwing than does C. eryphon (Brown, 1957; Clench, 1961; Hooper,
1973). These latter two characters show considerable individual vari-
ation as well as being dependent upon specimen condition, conse-
quently their usefulness for identification is limited and prone to sub-
jective interpretation.

Presence or absence of the mid-cell discal bar is perhaps the most
heavily relied upon distinguishing character used in keys and/or re-
gional works. However, no mention of variability of this character is
made.

Some specimens of C. eryphon possessed a distinct or variously
smeared mid-cell bar (d in Fig. 5). Of 41 specimens of C. eryphon
examined 7 (17%) possessed a mid-cell bar or spot and 5(12%) had
that area smeared by dark scaling thus negating determination. Geo-
graphic range of these specimens varied from British Columbia to the
Northwest Territories but most (36) were from Alberta.

The mid-cell bar, typically present in C. niphon was smeared or
rarely absent. Fifty specimens were examined (49 Alberta) and 3 (6%)
had the bar absent while 9 (18%) had that area smeared or obscured.
Extreme variation of other characters including wing checkering; an-
gle of fore- and hindwing chevrons; shape of median bands; hindwing
crennation; and most especially color of markings, distribution of gray
scaling and ground color was quite evident in the sample of Alberta
C. niphon. It is unknown whether such variation is typical of the
species in other parts of its range. Enumeration and description of
this variability of C. niphon is intended in a later paper.

In conclusion, when using available literature to key or identify

od

Moss, 1949). Question mark indicates an unconfirmed locality for C. eryphon. 2, Cal-
lophrys niphon clarki from Clyde, Alberta taken on 7 May 1978. x5, a, b, c explained
in text.

252 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

5

Fics. 3-5. 3, Callophrys eryphon eryphon from Seebe, Alberta taken on PAM
May 1978 (typical form with single discal cell bar). 4, Callophrys niphon clarki from
English River, Ontario taken on 17 February 1965. 5, Callophrys eryphon eryphon

from Terrace, British Columbia taken on 3 June 1960 (atypical form with two discal
cell bars). x5, a-d explained in text.

VOLUME 33, NUMBER 4 253

suspected specimens of C. eryphon or C. niphon care should be ex-
ercised. Comparison with long series of each species is desirable; and
recognition that character variability could potentially lead to erro-
neous identifications is necessary.

ACKNOWLEDGMENTS

I thank Dr. C. D. Bird (University of Calgary, Calgary, Alberta) and
Mr. J. D. Lafontaine (Biosystematics Research Institute, Ottawa) for
confirming the identification of C. niphon clarki. Mr. Lafontaine kind-
ly arranged for the loan of specimens from the Biosystematics Insti-
tute collection. Mr. T. W. Thormin pointed out the lack of Alberta
records for C. niphon and allowed me access to his collection. Dr.
Bird and Extension Entomology, University of Alberta allowed access
to collections in their care. Dr. Bird criticized early versions of the
manuscript and provided distribution records for Alberta C. eryphon.
Dr. D. Barr (Royal Ontario Museum, Toronto) criticized a later version
of the manuscript. The manuscript was typed by ROM staff.

LITERATURE CITED

Brown, F. M. 1957. Colorado butterflies. Denver Museum of Natural History, Denver,
368 p.

CLENCH, H. K. 1961. Tribe Theclini in Ehrlich, P. R. and A. H. Ehrlich. How to know
the Butterflies. Wm. C. Brown Co., Dubuque, Iowa, 262 p.

Hooper, R. R. 1973. The butterflies of Saskatchewan. Museum of Natural History,
Regina, Saskatchewan, 216 p.

HowE, W. H. 1975. The butterflies of North America. Doubleday and Co., Garden
City, New York, 633 p.

Moss, E. H. 1949. Natural pine hybrids in Alberta. Can. J. Research (c) 27: 218-229.

ADDENDUM

Additional specimens of Callophrys niphon clarki from Alberta have recently been
identified by Dr. C. D. Bird. These specimens represent a northward extension of the
known range within the province of approximately 400 miles and a first record for the
species from the Northwest Territories. The collection data are as follows:

a) Pine Lake, Wood Buffalo National Park, Alberta: 59°34’N, 112°15’W, | June 1974
(1 specimen); 6 June 1976 (1 specimen), Elsie Kuyt, lakeshore, sandy with scattered
willow and aspen poplar with white spruce, Banksian pine and mature aspen further
back from shore.

b) Fort Smith, Northwest Territories: 60°01’N, 111°52’W, 29 May 1976 (1 specimen),
Elsie Kuyt, domestic garden adjoining aspen poplar—whitespruce—Banksian pine,
mixed forest.

I am indebted to Elsie Kuyt for allowing me to use her collection data and Dr. Bird
for bringing these data to my attention.

Journal of the Lepidopterists’ Society
33(4), 1979, 254-257

LIFE HISTORY OBSERVATIONS ON HEMARIS GRACILIS
(SPHINGIDAE)

BENJAMIN D. WILLIAMS
The Lawrence Academy, Groton, Massachusetts 01450

ABSTRACT. The mature larva of Hemaris gracilis is described and figured from
material at Groton, Middlesex County, Massachusetts. The foodplant is low bush blue-
berry, Vaccinium vacillans Torrey.

According to recently published material (Hodges, 1971), the im-
mature stages of Hemaris gracilis are unknown. There is uncertainty
about the identity of its foodplant. This report describes the larval
stages and identifies a foodplant for this species.

On 27 May 1978 in Groton, Middlesex Co., Mass., I was looking for
Hemaris gracilis adults since for several years previously I had taken
individuals of this species feeding at the blossoms of early low bush
blueberry, Vaccinium vacillans. At approximately 1300 I observed a
female hovering over the V. vacillans but obviously not feeding. As
I observed her, she oviposited on the underside of new growth at the
extremity of a twig. The egg was retrieved; in the process I lost sight
of the adult. With the exception of its very small size the egg was
typically sphingifoGrm—pale green and slightly oblong in shape. The
color perfectly matched that of the leaf on which it was placed.

I returned to the same area on 30 May and captured a female of H.
gracilis which I subsequently placed in a “flying” cage. On 1 June
1978 a third female was observed ovipositing on V. vacillans at 1630;
I was able to retrieve four eggs before the female disappeared. In the
meantime I had put cut twigs of V. vacillans in the cage with the
female taken on 30 May. There were some blossoms on the vacillans;
in addition lilac blessoms were placed in the cage as a source of food
for the moth. Although I never actually saw the caged moth feed or
oviposit, she lived for five days. After the moth died I recovered twen-
ty eggs from the cut V. vacillans. The location of the eggs on the plant
was the same as it had been in the wild but the eggs tended to be
laid in small clusters rather than singly as was the case with the free
flying females.

A total of twenty-five eggs was collected from three females. The
larvae were reared through the fourth instar on V. vacillans which
was cut with the stems placed in water in the rearing cage. The fol-
lowing calendar was maintained on the first egg retrieved.

27 May egg retrieved

2 June larva hatched—typically sphingiform, pale green in color
with a black caudal horn

VOLUME 33, NUMBER 4 259

8 June larva entered second instar
12 June larva entered third instar; color patterns become apparent
19 June larva entered fourth instar
25 June larva entered fifth instar

1 July larva entered prepupal stage; body color takes on a purple

shade

2 July larva spins a loose cocoon in debris at the surface of the
ground (peat moss in this instance)

8 July pupation occurs; the pupa is active but not otherwise dis-
tinctive
4 August an adult Hemaris gracilis ¢ emerges; appears normal in all
respects

The larvae from the three gracilis females showed no significant
variation either among themselves or in color patterns throughout
their growth stages. Of the twenty-five larvae, twelve pupae were
obtained. Two larvae were preserved in alcohol, two died as a result
of accidental injury and the other nine died of unknown causes. The
larvae were transferred to high bush blueberry, Vaccinium corym-
bosum L.., for the fifth instar as I relocated for the month of July, and
V. vacillans was not available. Since no mortality occurred during
this instar, and the larvae readily accepted the foodplant, it is likely
that V. corymbosum is a host for Hemaris gracilis in areas where it
and the moth coexist.

Of the twelve pupae, four moths emerged in late summer—males
on 4 and 11 August, females on 6 and 10 August. The remaining pupae
are viable and will overwinter. It would appear that in New England
there is a partial second brood of Hemaris gracilis which flies in
August but for the most part the species is single brooded.

Description of the mature larva. Length in resting position 40-48 mm; width of
head 6.8 mm; height of head 6.6 mm. Basic color yellow-green. Head slightly darker
than body, dull, with extremely fine granulations; a narrow, dark brown line enclosing
the anterior four ocelli. Prothorax with a narrow, slightly raised anterior ridge (“cervical
shield’) extending down on each side to about twice the height of a spiracle above the
level of the dorsal end of each spiracle, light yellowish white, posteriorly tinged with
pink, shiny, with a few slight rugosities. Spiracles pink, each with a small white dot
in each dorsal and ventral end; sometimes with a small, orange-brown area anterior to
it. Entire dorsal area slightly darker green than remainder of body, unmarked, with
dorsal aorta showing through slightly darker. On each side a thin, yellowish white line
beginning at anterior edge of mesothorax and running uninterruptedly to base of caudal
horn. At their anterior ends these lines are slightly closer together than the width of
the head; gradually diverging to abdominal segment 2, thence parallel to last abdominal
segment, then converging to base of caudal horn. A short, yellowish white line along
each edge of tergite of posterior abdominal segment, extending to this segment’s point-

256 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fic. 1. Mature larva of Hemaris gracilis. A, showing dorsolateral yeiiowish-white
line. B, showing dark ventral surface. (Photographs by A. B. Klots.)

ed posterior end. Skin ef body smooth, sparsely and very minutely setose with fine,
white dots in 7-9 irregular, transverse rows on each segment. Ventral surface of body
dark reddish to purplish brown, this broken on mesothorax and metathorax, thence
continuous to anal prolegs, extending up on sides to above bases of prolegs, there
darker, and with a very slight yellowish upper edge. Legs pinkish brown, darkening
to tips. Prolegs of abdominal segments 3-6 very dark purplish brown; last prolegs
green, purplish brown ventrally. Horn light reddish brown, darkening toward tip, with
sparse setiferous rugosities, its terminal two thirds shinier; slightly shorter than dorsal
length of 8th abdominal segment.

Prepupal condition. Entire dorsal area suffused with dull reddish brown, extend-
ing laterad to include light dorsolateral lines. Head slightly duller green. Light, raised
prothoracic ridge, green lateral areas and dark ventral areas unchanged.

Characterization. From the larvae of H. thysbe (Fabricius) and diffinis (Boisduval)
(see Forbes, 1948, pp. 182-184 and 195-196) the larva is strongly differentiated by its
solidly dark ventral surface. Compared with thysbe the head and body are smoother,
the prothoracic shield narrower and smoother, the horn shorter and the dark, pale-
edged dorsal line absent. In these characters it seems to be more like diffinis.

Foodplant. Vaccinium vacillans Torrey is a preferred foodplant during the larval
stage of Hemaris gracilis. The strong probability exists that other species of Vaccinium
also serve as host plants for the moth throughout its range.

Disposition of specimens. The female which was originally caged is pinned and
in the author’s collection. Four moths emerged from pupae, 2 6 and 2 @. All, with the
eile of one female whose wings failed to expand, are pinned and in the author's
collection.

ACKNOWLEDGMENTS

I thank Dr. Alexander B. Klots of Putnam, Connecticut for his in-
valuable assistance in caring for the larvae during periods of my ab-

VOLUME 33, NUMBER 4 Zoe

sence, describing the mature larva and taking the photographs which
accompany this article. I also credit Dr. Hermann Flaschka, Chem-
istry Department, Georgia Tech, Atlanta, Georgia, for his advice as to
how to construct a flying cage in order to obtain eggs from the cap-
tured female.

LITERATURE CITED

FORBES, W. T. M. 1911. A structural study of the caterpillars II. The Sphingidae. Ann.
Entomol. Soc. America 4: 261-279.

1948. Lepidoptera of New York and neighboring states. Part I], Geometridae,
Sphingidae, Notodontidae, Lymantriidae. Mem. 274, Cornell Univ. Agric. Exper.
Sta., Ithaca, N.Y.

HODGES, RONALD W. 1971. Sphingoidea. Fasc. 21, The Moths of America north of
Mexico, p. 117. E. W. Classey Ltd., & R. B. D. Publications, Inc.

Journal of the Lepidopterists’ Society
33(4), 1979, 257

GENERAL NOTES

NEW PAPILIO CRESPHONTES HOSTPLANT

In mid-April 1978 I was examining Torchwood (Amyris elemifera L.) shrubs in the
understory of second-growth dry hammock on Big Pine Key, Monroe Co., Florida. No
rain had fallen for a month, and this particular place was exposed to a relentless parch-
ing southeast wind that had wilted all Torchwood at the hammock edge. But inside
the hammock, the shrubs looked healthy. Here I was surprised to find ova and larvae
in all instars of Papilio cresphontes cresphontes Cramer in circumstances theoretically
more suitable for P. aristodemus ponceanus Schaus. Furthermore, a few cresphontes
were flying through shaded hammocks here and in known Upper Keys ponceanus
habitats. Several cresphontes females investigated Torchwood but I witnessed no
oviposition. The few eggshells found were not necessarily on the youngest growth,
and first instar larvae accepted older growth. I gave one of these larvae new leaves
from Torchwood growing in full sunlight, and it ate them readily. When I tasted these
leaves they had a sharp tang almost like that of mint, followed by a longer-lasting bitter
aftertaste. Shade-grown new leaves lacked both these extremes.

I brought six final instar larvae back to New York, hoping to rear them through to
adults even though I had no Torchwood growing at home. Surprisingly, they refused
mature leaves of Citrus paradisi Macf. and etiolated shoots of Ruta graveolens L. After
wandering in the cage for some days, all pupated. Except for two partly abortive pupae
which I preserved, the rest emerged as characteristic but undersized adults.

FRANK RUTKOWSKI, 153 Centre St., New York, New York 10013.

258 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

Journal of the Lepidopterists’ Society
33(4), 1979, 258

ERYNNIS BAPTISIAE (HESPERIIDAE) ON CROWN VETCH (LEGUMINOSAE)

Crown vetch, Coronilla varia L., is a European perennial leguminous ground cover
introduced to North America after 1890. It has been extensively planted by the Penn-
sylvania highway authorities to control embankment erosion. This program began in
the mid-1930s and accelerated in the past 20 years (Wheeler, 1974, Can. Entomol. 106:
897-908). Crown vetch is now thoroughly naturalized in Pennsylvania as a common
component of old-field successional vegetation; it has also spread to adjacent south-
eastern New York and northern Maryland and Delaware. Wheeler (loc. cit.) conducted
an insect survey of the plant and found two butterflies breeding on it (counties un-
specified): Colias eurytheme Bdv. (Pieridae) and Erynnis baptisiae (Forbes) (Hesper-
iidae). He did not consider either to be of potential economic importance.

Shapiro (1966, Butterflies of the Delaware Valley, p. 53) judged E. baptisiae to be
“locally common” in southeastern Pennsylvania but did not find it on Coronilla, al-
though C. eurytheme was recorded on that plant (p. 38). In 1966 and 1967 it was noted |
as singletons in the vicinity of Coronilla in Montgomery and Chester Counties.

From 10-12 July 1978 I collected intensively in areas of Montgomery, Delaware,
and Chester Counties, Pennsylvania which I had often visited from 1955 through 1966.
In many of these localities I found E. baptisiae the commonest butterfly, a situation
never previously observed. Where Coronilla was abundant E. baptisiae usually out-
numbered all other butterflies and skippers combined. Numerous ovipositions on
crown vetch were observed, and, notably, male “territoriality’—which is very con-
spicuous in low-density populations of E. baptisiae—was much reduced or even ab-
sent. Thirty specimens were collected in an hour at a 0.5 ha stand of the plant in eel
Township, Delaware Co.

This unprecedented abundance may reflect an upward shift in the carrying capacity
of the environment for E. baptisiae in the presence of a newly adopted, exotic host. A
precisely parallel situation is believed to exist with populations of Pieris napi micro-
striata Comstock (Pieridae) on introduced watercress in California (Shapiro, 1975, J.
Res. Lepid. 14: 158-168). Ongoing monitoring of the range and population levels of E.
baptisiae where Coronilla occurs would be most desirable.

ARTHUR M. SHAPIRO, Department of Zoology, University of California, Davis, Cal-
ifornia 95616.

Journal of the Lepidopterists’ Society
33(4), 1979, 258-260

DOES HESPERIA JUBA (HESPERIIDAE) HIBERNATE AS AN ADULT?

The life history of Hesperia juba Scudder is very poorly known. The early stages
were described by MacNeill (1964, Univ. Calif. Publ. Entomol. 35: 67-77), who ob-
served that “the adults are present from April through October, with some variation
according to locality; evidently emergence is rather continuous and there are no distinct
seasonal broods.” This was inferred from the data on 769 specimens from hundreds of
localities, not incorporating long series from single places. For southern California,
imme! and Emmel (1973, Butterflies of Southern California, p. 84) record two broods,
April-June and August-September. This is closer to the picture which emerges when

VOLUME 33, NUMBER 4 259

long series from single localities in northern California are examined. Since 1972 reg-
ular, frequent butterfly sampling has been done in the Donner Pass vicinity (2,100 m),
Nevada and Placer Counties, as part of a larger phenological study. During this period
525 individuals of H. juba have been examined. Emmel and Emmel (1962, J. Lepid.
Soc. 16: 36) collected in Donner Pass from 17 June-26 August 1960 and recorded H.
juba from 17 June-1 July only. My records, given below, corroborate this spring flight
but also indicate an autumn flight missed by the Emmels:

Spring
Year Locality Spring dates Fall dates snowpack
1972 Boreal Ridge v. 24-vi. 7 viii. 10-x. 4 light
1973 Soda Springs-Norden not seen Xs key heavy
1974 y vi. 9 viii. 24—ix. 27 heavy
1975 H Witsoe he Pra, SO) moderate
1976 v. 14-vii. 1 viii. 20-x. 8 very light
1977 i vi. 4 ix. 2-ix. 23 light
1978 é vi. 14-vii. 1 Vint. L5=x.23 very heavy
1979 u vi. l-vii. 12 ix. 4—ix. 30 moderate

This is an unusual phenology for a skipper, especially in montane habitats. The fall
flight appears just as rabbitbrush (Chrysothamnus nauseosus (Pall.) Britton, Compos-
itae) comes into flower and ends just as the last individuals of that species go to seed.
During this time nightly low temperatures are usually near to below 0°C. Twenty or
more H. juba may be found on individual rabbitbrush plants, and they are rarely seen
elsewhere: feeding occurs throughout the day if the sun is shining and the air warm.
Many other insects also visit Rabbitbrush at this time, including tachinid and syrphid
flies, a wide variety of bees including Bombus spp., several day-flying noctuid moths,
and such butterflies as Vanessa virginiensis Drury and Polygonia zephyrus Edwards
(Nymphalidae), Apodemia mormo Felder & Felder (Riodinidae), Neophasia menapia
Felder & Felder (Pieridae), and Ochlodes sylvanoides Boisduval (Hesperiidae). All of
the autumn H. juba are fresh, with bright green ventral hindwings bearing lustrous
silvery spots; only at the very end of the flight, in October, do any individuals show
noticeable wear or fading, and this is uncommon.

In spring H. juba appears shortly after snowmelt, often flying on warm south slopes
while the north-facing ones are under 2 m of snow. At this time most flower visits are
to dandelion (Taraxacum officinale L., Compositae) but many individuals can be seen
“body basking” on the bare ground in sunlight, courting, and ovipositing. The popu-
lation density in spring is consistently about one-tenth of the previous autumn’s levels,
and contrary to MacNeill’s observations there has been little year-to-year variability in
numbers. The spring flight is very short, often observed only in one weekly sample.
In 1976, a year of record low snowpack, H. juba was fairly common on 14 May and
singletons were seen weekly to 1 July.

Most strikingly, the average condition of spring individuals is much poorer than in
autumn. Although a few could be called “fresh” in appearance, most have the green
faded to dull brown and the silver entirely lost. Under a dissecting microscope such
specimens show up to 20% scale loss on the ventral hindwing. Dorsally the golden
ground color is usually intact but the dark borders appear somewhat faded.

All this points to adult hibernation. Of the five species of butterflies listed above as
visiting rabbitbrush, the two nymphalids reappear at the same time as H. juba and are
generally considered to be hibernators. Like H. juba they are very fresh in autumn but
appear relatively worn in spring. Also like H. juba, they feed almost continuously from
rabbitbrush in good weather. The other three species do not hibernate as adults and
have only an autumn flight. They all feed only at midday to mid-afternoon. Adult
hibernation has never been reported in the family Hesperiidae in rigorous climates,
but the circumstantial case for it at Donner Pass is duplicated by admittedly less mas-
sive documentation in Trinity, Sierra, Plumas, Siskiyou, Shasta, and Alpine Counties.

260 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

In the Great Basin at lower elevations H. juba also flies at rabbitbrush season and
engages in massive, continuous flower visitation.

Could a spring generation emerge in June from eggs laid the past September at
Donner? It is difficult to see how. Skippers grow slowly as a rule; in optimal weather
development from a June egg to a fall adult requires 8-11 weeks. A September egg
would have, on the average, 6 to 8 weeks before continuous snow cover developed.
During this time the days are shortening rapidly, and ambient temperatures exceed
10°C for only a few hours a day while nightly minima may reach —10°. An increasing
number of days have little or no sunshine. MacNeill found that captive larvae kept
outdoors in the San Francisco Bay Area would not feed in cool, cloudy weather but
resumed activity immediately in strong sunshine or at indoor temperatures. It is dif-
ficult to envision much activity or growth after early October at Donner, and impossible
to envision under snow. Moreover, the grasses are in poor condition at this time. The
most important spring host seems to be Agrostis idahoensis Nash, which is mostly
brown in October but grows rapidly after snowmelt. After snow leaves an area adults
are generally not seen for 2 or 3 weeks. Ambient temperatures are often high and the
days are quite long, but this is a very short developmental time for a skipper! Moreover,
most of the H. juba at Donner in spring are worn when first encountered. If little or
no wear is shown by fall adults three weeks into the flight, why should newly emerged
spring animals deteriorate so rapidly?

It is not inconceivable that H. juba could have a “mixed strategy” of overwintering
as adults, eggs, and perhaps young larvae. The two latter stages are common among
Hesperiines. Absolute proof of overwintering will require detection of hibernating
adults in midwinter, or the recapture in spring of individuals marked the previous
autumn. In the absence of such direct evidence the inferential case is rather compel-
ling.

ARTHUR M. SHAPIRO, Department of Zoology, University of California, Davis, Cal-
ifornia 95616.

Journal of the Lepidopterists’ Society
33(4), 1979, 260-261

A MIGRATORY FLIGHT OF URANIA FULGENS (WALKER) IN HONDURAS
(URANIIDAE)

Urania fulgens has long been known as a migratory diurnal moth. Williams (1930,
The Migration of Butterflies, Edinburgh; 1958, Insect Migration, London) has provided
excellent summaries of earlier published observations on their migratory flights. These
include records from most of Central America from Mexico to Panama and in Colombia
and Ecuador. Although we have seen no prior records of flights in Honduras (they may
exist), there was no reason not to expect them there.

Williams (1958, op. cit.) tabulated over sixty records of flights from Central America
and found in these some evidence of a change of direction of flight at different seasons
(including return flights from Mexico and Costa Rica). He found reports of flights from

March to August, with a preponderance to the north in March and April and to the east
or southeast in June and September. However, he concluded that more records and
more accurate compass-directions are needed. On this basis the following brief note
is offered.

On 24 August 1978 while seated on a terrace of the Gran Hotel Tela, Tela, Distrito
Atlantida, Honduras, during a light rain near mid-day, we noticed a number of Urania

VOLUME 33, NUMBER 4 261

fulgens (Walk.) in flight at an elevation of 10-12 m above ground level. The flight
pattern was not random, but directional, and generally southwesterly around two sides
of the terrace. The following morning at 0600, upon walking onto the front balcony of
the hotel which faced on the main street of Tela, we saw large numbers of individuals
flying along the street from northeast to southwest. They flew singly or in small groups
at levels ranging from one or two m to 12 or 15 m above the street level. The sky was
overcast and a light rain was falling after intermittent heavy tropical showers during
the night. Five minute counts yielded more than 150 individuals and we estimate that
1,500-1,800 flew past during the hour in which they were under observation. When
we terminated observation at 0700, there had been no visible let-up in the flight.

At about 0800 we left Tela by car for El Progresso. This section of highway runs in
a generally northwesterly direction and in the northwesterly stretches, the moths con-
tinued to cross the highway toward the southwest in large numbers over a distance of
at least 30 km but began to disappear or lose their directional flight as the road moved
into the mountains.

Where the road crossed the flight lanes, hundreds of dead or stunned moths were on
the highway. Stopping to collect a sample of perfect specimens we found that ants
reached them within minutes, eating the abdomens of still-living individuals. However,
we obtained 39 freshly-emerged specimens in satisfactory condition in half a dozen
brief stops. The sex ratio in our samples was 26 66:13 2Q.

Thus, in the flight observed by us in late August, the direction was from northeast
to southwest, in contrast to reports of others for this general time period.

JOHN A. CHEMSAK, E. G. LINSLEY & JUANITA M. LINSLEY, Division of Entomology
and Parasitology, 201 Wellman Hall, University of California, Berkeley, California
94720.

Journal of the Lepidopterists’ Society
33(4), 1979, 261-264

CAPTURES OF LARGE MOTHS BY AN ULTRAVIOLET LIGHT TRAP

Early in April 1978, J. Muller installed a standard black light trap, made by the
Ellisco Company, on C. B. Worth’s farm in Eldora, Cape May Co., New Jersey (Fig.
1). This trap, plugged into an ordinary electric outlet, uses a tube of only 15 watts,
emitting both visible blue and ultraviolet (black) rays. Insects striking the four vertical
baffles surrounding the tube fall through a funnel into a collecting chamber containing
cyanide or other lethal volatile chemicals. The trap is standard equipment for agents
of Rutgers University monitoring the abundance of flying phototropic agricultural pests
throughout the state of New Jersey.

Muller had been collecting moths on this farm by other means since August 1972,
as part of his statewide survey of the macrolepidoptera of New Jersey (Muller, 1976,
J. New York Entomol. Soc. 84: 197-200, and unpublished). Since the farm is isolated
and not close to competing lights, this new trap presented an opportunity to study the
extent to which an ultraviolet light diverts moths from their natural nocturnal functions.
It was decided to record all sphingids and larger saturniids that were removed from
the environment at this focus (Table 1).

For a few days, from time to time, the trap did not operate well because of exhaustion
of the lethal gases. However, the kill (summarized in the table) remains representative
for comparative purposes, since the flight period of most species occupies several
weeks. In the case of double-brooded species there must obviously be two peaks of
abundance; these have not been separated in the table.

262 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

Fic. 1. Standard black light trap, the Ellisco Co.

The grand total of moths killed, 330, may look impressive, but compared to the
number of smaller forms taken it is insignificant. Each night’s catch contained perhaps
as many as a half dozen large moths, along with a pint to a quart of smaller ones. The
latter fraction must have numbered several hundreds. Thus, over a period of about four
months, the trap caught in the neighborhood of 50,000 moths. This may have repre-
sented the removal of a considerable number of ovipositing females, and thereby ought
to have affected the biomass of foraging caterpillars in the population of 1979.

However, it is difficult to draw conclusions when the status of captured moths is
unknown. Have females already laid their eggs? Have males already mated? If both
the answers are “Yes,” removal of the moths would not affect next year’s crop.

ixamining the catches of the four commonest species (in those cases when the sexes
were largely known), we see that among luna, polyphemus, io, and imperialis, 138
males and 13 females were taken (91.4 percent males and 8.6 percent females). These
figures suggest first that females may be less mobile and secondly that they may be
more strongly motivated to fly on ovipositional errands rather than be diverted by
attractive lights.

VOLUME 33, NUMBER 4 263

TABLE 1. Large moths caught in a black light at Eldora, New Jersey, 2 May to 12
September 1978.

Sex
Species M F ? Total Inclusive dates
Actias luna (Linnaeus) ays, Al 2 DA ee? V to 2 1X
Deidamia inscripta (Harris) 1 1 7 Ona VatowlG) Vil
Antheraea polyphemus (Cramer) Pas fr SEAS WY GoeVeto S VILL
Paonias astylus (Drury) Oo 8. 23 BY AOW to 22 0
Smerinthus jamaicensis (Drury) —_ — 2 D2 NW to JA Wal
Lapara coniferarum (J. E. Smith) ho A AAS) DSW Teo) WAU
Hyalophora cecropia (Linnaeus) 3 0—- — ® COW io IW
Paonias myops (J. E. Smith) —_—_ — D 230) Veto 29) Vir
Darapsa pholus (Cramer) Ley alt LG WS SOW ww NH Id
Cressonia juglandis (J. E. Smith) t 4 5 30V to 5 VIII
Callosamia promethea (Drury) a 8 LW t JEW
Paonias exaecatus (J. E. Smith) 4 dh i DY aL AL toy DE WAU
Sphecodina abbottii (Swainson) —— 1 eval
Amphion nessus (Cramer) —_- — 1 he Ab AW
Automeris io (Fabricius) Ses Lo 85 4 Wii sO inl
Sphinx gordius Cramer Gs 3 9 12 VI to 30 VI
Ceratomia catalpae (Boisduval) —_— — 3 S 1d Wl iw & IDX
Manduca quinquemaculata (Haworth) —— 3 4 26 VI to 24 VIII
Eacles imperialis (Drury) BU HB = BY sO Wo 2 in
Citheronia regalis (Fabricius) ine ea — YF WAU ww BS WAU
Hyles lineata (Fabricius) 1 — 1 D2 XS NVA® A WAU
Dolba hyloeus (Drury) —_ — IL ih HAO
Eumorpha pandorus (Hubner) —- — ] ik SQ WADE
Paratrea plebeja (Fabricius) —_—- — 1 1 24 VIII
Manduca sexta (Linnaeus) a 2 DY SIL WVU to © IDK
Total Kon) Zier TSO 330

The obvious question then becomes whether or not males are so drawn to lights that
they do not respond to female pheromones. One test of that possibility gave inconclu-
sive results. During the course of these observations, Worth tethered 16 newly emerged
Citheronia regalis females within a few hundred yards of the light trap. They were
definitely in competition with the ultraviolet light source, and cruising wild males had
an easy choice of which target they would select. Fourteen females secured wild mates,
while the trap took only one male. However, during the same three-week interval, 15
reared and marked males were liberated but none of these was trapped. This species
is apparently only mildly phototropic.

A further suggestive finding was that the two trapped female Eacles imperialis that
were dissected were found to be devoid of eggs.

As an incidental observation, it is interesting that Callosamia promethea, a common
species in this region, was represented in the trap by three females but no males, the
latter being largely diurnal.

This study bears on the question of rather new “light pollution” as it relates to
populations of phototropic insects. For several decades it was presumed that new in-
secticides such as DDT were responsible for the decline of large moths in our great
urban centers and their suburbs. However, it has not been clear in more recent years
why these insects survived in regions such as Worth’s farm in Eldora, New Jersey,
where DDT and related insecticides have been used vigorously to combat agricultural
pests, mosquitoes and gypsy moths.

264 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

The difference may lie in rapidly increasing popularity of mercury vapor lamps for
urban street lighting as well as for community and private use. These emit ultraviolet
light. Given the number of these light sources, insects must be attracted in inestimable
numbers, perhaps withdrawing them from reproductive duties to the point of local
species extinction. Of course such lamps do not kill insects, but they immobilize them,
rendering them as biologically inactive.

In rural areas the use of this type of illumination is much less common. This may
account for the greater abundance of large moths in these areas.

Finally, the light trap contained many other orders of insects, among which Hyme-
noptera were abundantly represented. None of these was saved for identification, but
the possibility remains that some were parasitoids of large moths. In such a case black
lights might have a favorable effect on moth populations.

C. BROOKE WoRTH, Eldora, R. D. Delmont, New Jersey 08314, AND JOSEPH MULLER,
Rt. #1, Lebanon, New Jersey 08833.

Journal of the Lepidopterists’ Society
33(4), 1979, 264-265

JAMES GRAHAM COOPER (1830-1902)

James Graham Cooper was an important 19th century naturalist in California. Col-
lections of Lepidoptera made by him became the basis for several new species de-
scribed by H. H. Behr, including Melitaea quino. Other entomological material he
collected was the basis for new taxa described by J. L. LeConte (Coleoptera) and J. W.
Greene (Hymenoptera). New species that were named after him include Anthocaris
cooperii Behr, Melitaea cooperi Behr, both Lepidoptera; and Lytta cooperi LeConte,
and Amphicoma cooperi (Horn), both Coleoptera. A very brief biography by Essig
(1931, A History of Entomology, Macmillan, New York) is sketchy and inaccurate. More
complete biographies by Grinnell (1905, The Condor, Vol. 5) and Emerson (1899, Bull.
Cooper Ornithol. Cb., Vol. 1) are more complete, but only cite his achievements in
ornithology. In researching some of the species of Lepidoptera named by Behr, I have
uncovered a fair amount of information on Cooper that may be of benefit to other
lepidopterists in the future.

James G. Cooper was born in New York City 19 June 1830. His father was a close
friend of James Audubon. He had an early interest in natural history and in 1850
accompanied LeConte on a collecting trip to California. After graduating from the
College of Physicians and Surgeons in New York in 1853, he took a position as phy-
sician and naturalist on an expedition exploring a potential railroad route through Or-
egon. In 1861 he was back in California and petitioned J. D. Whitney to join Whitney's
California Geological Survey as zoologist. For the next several years he did work with
Whitney off and on with the Survey Team. Whitney’s chief assistant, W. H. Brewer
described him as “a man of more than ordinary intellect and zeal in science, but not
a very companionable fellow in camp” (1966, Up and Down California, Univ. California
Pr., Berkeley). His primary duties with the Survey were to collect plant specimens, but
his primary interest was vertebrate animals and not botany as was cited by Essig (op.
cit.). During the 1860’s he became associated with Behr and the California Academy

of Science in San Francisco. His primary interests during this period were fish (both
marine and freshwater) and marine animals, and he presented many papers to the
Academy describing new species. During this period he collected entomological ma-
terials that he supplied to Behr and other specialists. His explorations ended in 1866
when he married Rosa M. Wells of Oakland. He practiced medicine in Oakland, where

he lived until 1871. In 1871 he moved his practice to Ventura County and his close

VOLUME 33, NUMBER 4 265

association with the California Academy of Science ended. In 1875 he moved to Hay-
ward and continued to practice medicine there until his death in 1902. He continued
his studies in Natural History during this time and The Cooper Omithological Club,
named in his honor, was organized in 1893.

JOHN H. MASTERS, 25711 North Vista Fairways Drive, Valencia, California 91355.

Journal of the Lepidopterists’ Society
33(4), 1979, 265

A RECENT RECORD OF SPEYERIA IDALIA (NYMPHALIDAE)
FROM MANITOBA

On 20 July 1977, Brook Nero (546 Coventry Road, Winnipeg, Manitoba) captured a
specimen of Speyeria idalia (Drury) in a prairie field, beside Assiniboine Forest,
Charleswood, Manitoba. The specimen is a male with a wingspan of 8.7 cm, and it is
not too worn. Assiniboine Forest is a 700 acre tract set aside as a natural park by the
city of Winnipeg. It is primarily an area of second growth aspen and oak. The collecting
site lies within the Park on the west edge, and has been identified as a potential
reclamation area to the original prairie. At present, however, it is largely bluegrass with
only a dozen or so surviving prairie forbs.

This is the only recent record for S. idalia in Manitoba. G. S. Brooks (1942, Canad.
Entomol. 74: 31-36) recorded a previous record from “Winnipeg” with the comment
that it was a stray that “almost certainly does not breed in the province.” It is unlikely,
however, that either of these Manitoba records represent strays. More likely they are
evidence of small colonies of the species still persisting on tiny remnants of virgin
prairie. The larval foodplant of S. idalia, the birds-foot violet (Viola pedatifida) is an
obligate species of mesic prairies, and adult butterflies seldom stray far from areas
where it grows.

The two Manitoba records represent the most northerly known records for S. idalia.
However, the species may have been widespread in occurrence on virgin prairie all
across southern Manitoba before these prairies were plowed and converted to wheat

fields.

JOHN H. MASTERS, 25711 North Vista Fairways Drive, Valencia, California 91355.

Journal of the Lepidopterists’ Society
33(4), 1979, 265-266

ABERRANT SPECIMEN OF LYCAEIDES MELISSA MELISSA
(LYCAENIDAE)

The accompanying photo (Fig. 1) shows the ventral view of two fresh specimens I
caught while collecting along the road to Deer Creek Campground, west of Heber City
(Wasatch Co.), Utah 23 June 1976. The specimen on the right is a normal female
Lycaeides melissa melissa (W. H. Edwards); the one on the left represents an aberration
in which the postbasal spots are lacking and the postmedian spots are almost lacking.
The extremely well developed marginal band of crescents indicates that the specimen
is referrable to Lycaeides melissa melissa rather than to L. melissa annetta (W. H.
Edwards).

266 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fic. 1. A, aberrant female Lycaeides melissa melissa, ventral surface. B, normal L.
m. melissa female.

I have shown pictures of this aberration to Dr. Cyril F. dos Passos, Dr. John C.
Downey, Harry K. Clench and several other knowledgeable lepidopterists and none
can recall seeing a similar specimen, which suggests that such an aberration is rare in
the species melissa. The aberrant specimen and several normal specimens from the

same collecting site have been donated to the Allyn Museum of Entomology, Sarasota,
Florida.

WILLIAM B. WRIGHT, JR., 18 Clinton Place, Woodcliff Lake, New Jersey 07675.

Journal of the Lepidopterists’ Society
33(4), 1979, 266-267

POPULATION OUTBREAK OF CATOCALA PALAEOGAMA (NOCTUIDAE)

Early in the afternoon of 22 July 1978, I encountered large numbers of the oldwife
underwing moth, Catocala palaeogama Guenée at Illinois Beach State Park in NE
Illinois. The park, located on Lake Michigan approximately 1 mi E of Zion, Lake Co.,
provides an excellent wildlife habitat. Natural features of this extensive tract include
prairie, forest, marsh, dunes, stream and several miles of Lake Michigan shore. Low
dunes along the beach support an unusual assemblage of plants, including various
dune-associated grasses, bearberry (Arctostaphylos Uva-ursi) and trailing juniper (Ju-
niperus sibirica). Black oak (Quercus velutina), sand cherry (Prunus pumila), willow
(Salix sp.), New Jersey tea (Ceanothus pubescens), shrubby cinquefoil (Potentilla fru-
ticosa), prickly-pear cactus (Opuntia polyacantha), wild indigo (Baptisia tinctoria),
lead plant (Amorpha fruticosa) and a proliferation of other plants cover dunes further
ee attracting a variety of unusual moth and butterfly species from early spring until
ate fall.

The concentration of Catocala palaeogama was discovered on trunks of black oak
at a sandy picnic area close to the lake and adjacent dunes inland. ! first observed the
moths when I startled individuals at their resting sites, causing them to fly to other
trees nearby. From one to as many as five recently emerged specimens were found
roosting on nearly every sizeable tree, always on the shady side, from two to five feet

VOLUME 33, NUMBER 4 267

above the ground. When sunlight touched resting sites, individuals moved around the
trunk to shade. Observing and collecting was facilitated in these moths as they rested
on the same side of all trees at a given time. Forewing markings, showing the consid-
erable variation common in many Catocala species, blended well with the grey bark
of the oaks.

It would be difficult to estimate the number of underwing moths in this aggregation,
or to know how extensive the population was. But certainly many hundreds of speci-
mens were congregated within the park that afternoon. It is likely this phenomenon
occurred in this area in other years as well, but this was my first observation of such
a remarkable event. While C. palaeogama was the predominate species represented,
single specimens of C. lacrymosa Guenée and C. amica Hubner were collected. A
large series of C. palaeogama was taken, and a number of these, deposited in the
collection of Mr. Bryant Mather, Clinton, Mississippi, were subsequently positively
identified by Mr. Eric Quinter, American Museum of Natural History. Specimens of
the same catch were also deposited in the collections of Dr. Clifford D. Ferris (Laramie,
Wyoming), Mr. Patrick J. Conway (Downers Grove, Illinois), Mr. Mogens C. Nielsen
(Lansing, Michigan), and the Illinois Natural History Survey (Urbana, Illinois).

It is interesting to note that a collecting trip during the following weekend to the
same locality yielded no additional specimens of C. palaeogama after the supera-
bundance of the previous week. The area which had been alive with activity at that
time was now dead, so far as that species was concerned. However, specimens of other
Catocala species were collected during the second visit.

IRWIN LEEUW, 1219 Crystal Lake Road, Cary, Illinois 60013.

Journal of the Lepidopterists’ Society
33(4), 1979, 267-268

NOTES ON THE BIOLOGY OF BATTUS PHILENOR (PAPILIONIDAE)
IN CENTRE COUNTY, PENNSYLVANIA

Centre County occupies parts of two geographic provinces. The northwestern third
lies in the Allegheny Plateau Province, while the southeastern two thirds belongs to
the Ridge and Valley Province (Westerfeld, 1959, Pa. Agr. Exp. Sta. Bull. 647: 6-17).
The configuration of land surface in the latter is due to the folding of the rocks into
parallel mountain chains. These sandstone ridges are oriented from southwest to north-
east and average from 550-730 m in elevation. Separated by these ridges are limestone
valleys averaging about 310 m above sea level. Battus philenor (L.) is largely confined
to the stream trenches of these valleys. One such area is along Spring Creek adjacent
to the Benner Springs Fish Hatchery about 8.5 km northeast of State College. A small
but rather stable population has existed here since at least 1974.

B. philenor is bivoltine with flight periods from 13 May to 16 June and 6 July to 26
August. Fresh adults occasionally observed in September and early October represent
a partial third brood. However, these individuals, mostly males, are probably lost to
the population since it is doubtful that their progeny would have sufficient time to
reach the pupal (overwintering) stage prior to the onset of cold weather.

Males are frequently observed visiting mud puddles, or flying rapidly along the creek
and woodland trails and in adjacent meadows. Females are more secretive and are best
sought along woodland trails or in open woods. Both sexes prefer pink to purplish
flowers such as Hesperis matronalis L., Dipsacus sylvestris Huds., and various thistles
(Cirsium spp.). Associated butterflies of special note include Asterocampa clyton
(Boisduval and LeConte), Calephelis borealis (Grote and Robinson), and Erynnis lu-
cilius (Scudder and Burgess).

268 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

One of the reported foodplants of B. philenor, Asarum canadense L. (Howe, 1975,
The Butterflies of North America: 390), is locally abundant on moist, rocky hillsides
along Spring Creek. It was assumed that the larvae were utilizing this resource, al-
though none could be located. A similar observation is made by Harris (1972, Butter-
flies of Georgia: 158) for Bibb County where larvae used only cultivated Aristolochia
even though Asarum was present. On 2 June 1978, a female was confined with potted
A. canadense. She died after 10 days without oviposition. Internal examination re-
vealed that she was mated and had numerous mature ova. We then began a more
intensive search for the specific host.

On 4 August at about 1700 h, a female B. philenor was observed ovipositing on
Aristolochia serpentaria L. This is the only Aristolochia species in the county and is
rare and confined to the Ridge and Valley Province (Westerfeld, 1961, Castanea 26:
34). Additional small plants (about 30 cm in height) were subsequently located growing
singly or in small groups on the rocky, open-wooded hillsides adjacent to the creek.
Numerous ova and first to fourth instar larvae were found on these plants. Most ova
were laid on the petioles and/or margins of the upper leaves with up to four individuals
per plant. Plants growing in moist, brushy, shaded areas were larger and more luxu-
riant; however, ova and/or larvae were present in significantly lower numbers. A female
captured on 8 August and confined with potted A. serpentaria oviposited within hours
of confinement. The resulting ova plus 11 field-collected larvae were reared to pupa
using potted plants. Enormous quantities of these plants were required to complete
larval development and several field trips were needed to collect additional food. On
one of these trips several late-instar larvae were observed wandering apparently in
search of food. Ehle (1951, Lepid. News 5: 103) noted for Lancaster Co., Pennsylvania
that the required food (A. serpentaria) far exceeded the quantity available at the orig-
inal site. The last instar larvae consumed the leaves, seed capsules, and stems to within
about 5 cm above ground level. Several first instar larvae were transferred to A. ca-
nadense immediately after emergence and all died without eating. In addition, last
instar larvae temporarily confined with this species refused to eat. Saunders (1932,
Butterflies of the Alleghany State Park: 234) remarked that he was able to get B. phi-
lenor larvae to eat only Aristolochia and not Asarum except for one larva which “ate
a little Wild Ginger, but did not seem to like it.” Of the 25 larvae reared to pupa, 60%
diapaused and were refrigerated.

Conclusions which may be drawn from these observations are as follows: The use
of Asarum canadense as a larval food source is to be seriously questioned if not com-
pletely discounted. At the Spring Creek site, the amount, distribution, and availability
of the host plant, Aristolochia serpentaria, appears to be a significant factor in regu-
lating population size. Selection pressure toward producing adult females and larvae
with maximum search capabilities would be necessary to maintain a stable population
in the absence of immigration. Larval parasitism (or predation) was not investigated,
however, all of the field collected larvae (third and fourth instars) were successfully
reared to pupae.

FRANK D. FEE, 308 S. Corl St., State College, Pennsylvania 16801 and RICHARD
BOSCOE, 150 Ridge Pike, Apt. 201, Lafayette Hill, Pennsylvania 19444.

Journal of the Lepidopterists’ Society
33(4), 1979, 268-269

BOOK REVIEW

BUTTERFLIES OF SOUTH AUSTRALIA, by Robert H. Fisher, 1978. One of the series
“Handbooks of the Flora and Fauna of South Australia,” issued by the Handbooks
Committee for the South Australian Government, 8vo, soft cover, [iii] + 272 pp., 83

text figures, 16 color plates. Price $9.50 Australian (approximately $11.00 U.S.).

VOLUME 33, NUMBER 4 269

Beyond the above information, the book gives no clue as to where one might obtain
a copy. Word from Entomological Reprint Specialists is that they will stock it eventually
but have no copies at this writing.

This little carp aside, the book is a gem. The front matter includes a history of the
study of South Australian butterflies, short but informative explanations of classifica-
tion, life histories (and how to study and record them), anatomy, distribution, how to
make and keep a collection. In the back of the book is a systematic list of larval food-
plants and the butterflies that use them, a lengthy (6 pp.) bibliography, a lengthy
glossary (8 pp.), and an index.

The bulk of the book, of course, comprises the species accounts. The butterfly fauna
of Australia numbers in all 366 species of butterflies, of which 64 occur in South Aus-
tralia. Each of them is figured in color, generally both sexes and both surfaces (with
full data appended for each figured individual), and thoroughly discussed: references,
terse description, larval foodplants, life history, habits, distribution, abundance, seasons
of flight. The color photographs of adults are all variously reduced (% to %), which
does not diminish their usefulness except in the smaller species, particularly some of
the Hesperiidae and Lycaenidae. So much white space surrounds the figures of the
latter groups that they easily could have been expanded to life size at no extra cost,
and that is unfortunate.

The most outstanding features of this volume, however, are the beautiful black and
white illustrations of the living early stages. The photographs are clear and crisp, and
unbelievably numerous. As a rough estimate, about 80% of the species are so illus-
trated, some with supplemental color photographs as well. The figures for each species
usually include the egg, larva (often both young and mature), and pupa, in most in-
stances the first published illustrations of them.

The Butterflies of South Australia is clearly aimed at the local collector, who is
blessed thereby with a guide that lepidopterists in most other parts of the world would
envy: authoritative, detailed, packed with information and good illustrations, stimulat-
ing in its frequent mention of subjects still in need of careful study. Mr. Fisher knows
and presents his subject well, and I recommend his book heartily to butterfly students,
not just in South Australia but wherever they may be, if their interests extend even a
little beyond the parochial and the philatelic.

HARRY K. CLENCH, Section of Insects, Carnegie Museum of Natural History, Pitts-
burgh, Pennsylvania, 15213.

Journal of the Lepidopterists’ Society
33(4), 1979, 269-270

BOOK REVIEW

THE BUTTERFLIES OF ORANGE COUNTY, CALIFORNIA, by Larry J. Orsak, 1977. Center
for Pathobiology, Museum of Systematic Biology, University of California, Irvine. 349
pp., 7 halftone plates, 56 text figs., 4 maps. Paperback. Price: $4.00.

This book is more than a regional checklist; it is a treatise on butterflies designed to
be independently useful to the beginning collector. The species accounts which form
its main body and purpose are preceded by extensive introductory material on classi-
fication, variation, structure and behavior; much of it common to most butterfly man-
uals, but other aspects (e.g., sound production, hilltopping, nectaring) seldom treated
outside the periodical literature and consequently less easily available. Several appen-
dices contain additional general information.

In content, therefore, the work is exhaustive and informative. Its organization, how-
ever, is another matter. In this respect it suffers certain shortcomings which make its
use difficult, especially for one not already familiar with the fauna it covers.

270 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

The author evidently enjoyed the luxury of unlimited space. He quotes full label
data from Orange County material in several private and institutional collections. Large
blocks of these data from the same localities differ only in terms of collection dates and
number and sexes of specimens, with the localities repeated in line after line. Surely
this could have been condensed with no loss of information.

Text figures, many of them very useful ones of larval foodplants, are often inserted
into species accounts with their explanations in the same type face as the rest of the
text, so that its smooth flow is interrupted and one must search for the continuation of
the main text. This confusing feature could have been eliminated by the use of a
smaller type for the figure legends.

But by far the biggest problem is that the genera within each family are arranged
alphabetically instead of in scientific order, not only in the main text but in the ap-
pended checklist as well. I am entirely unable to comprehend the rationale for this
procedure. Thus, in the Lycaenidae one finds blues and hairstreaks jumbled together
with the coppers in their midst, while among the nymphalids such closely related
genera as Cynthia and Vanessa are separated by no fewer than seven others of more
distant affinities. The result is a feeling of complete disorientation, making an overall
impression of the area’s butterfly fauna at a glance very difficult to obtain.

Despite these reservations, all but the last comparatively minor, the book is a mine
of information if one is willing to do a little digging, modestly priced and a worthy
addition to the literature of the still-rich butterfly fauna of southern California. The
author's expressed primary purpose of recording distributions “before habitats are de-
stroyed and the memories of veteran Orange County collectors fade” is very com-
mendable and has been well realized.

RODERICK R. IRWIN, Illinois Natural History Survey, Urbana, Illinois 61801.

Journal of the Lepidopterists’ Society

33(4), 1979, 271-280

EDITORS’ NOTE

The editors wish to extend their sincere appreciation to all persons who have given
generously of both their time and knowledge while serving as reviewers for manu-
scripts submitted to the Journal during the past two years. The following persons have
reviewed one or more manuscripts during the preparation of Vols. 32 (1978) and 33

(1979):

William A. Anderson
Richard A. Arnold
Jose L. Barata

M. Deane Bowers
André M. Blanchard
Brian P. Bradley
Lincoln P. Brower
F. Martin Brown
Keith S. Brown, Jr.
Robert S. Bryant
Frances S. Chew
Cyril A. Clarke

J. F. Gates Clarke
Harry K. Clench
Don R. Davis

Jane V. Z. Dingman
Gerrit deBoer

John C. Downey
Joseph F. Doyle, III
Thomas C. Emmel
John H. Fales

Douglas C. Ferguson
Clifford D. Ferris
Richard S. Fisher
John G. Franclemont
Richard S. Funk
Lawrence F. Gall

Richard C. Gethmann

George L. Godfrey
Frank E. Hanson
D. F. Hardwick

S. J. Harrison
Ronald R. Hodges
Charles C. Horton
Rod R. Irwin

Kurt Johnson

Roy O. Kendall
Steven J. Kohler
John Lane

Don MacNeill
Lee D. Miller
Thomas A. Miller

Robert T. Mitchell
Raymond W. Neck
Herbert H. Neunzig
Paul A. Opler
Richard S. Piegler
Kenelm W. Philip
Austin P. Platt
George W. Rawson
J. C. E. Riotte
Theodore D. Sargent
Arthur M. Shapiro
Nancy E. Stamp
Ray E. Stanford
Fred W. Stehr
James G. Sternberg
John R. G. Turner
Paul M. Tuskes
David H. Wise
Allen M. Young

INDEX TO VOLUME 33 (Exclusive of Supplement)

(New names in boldface)

Amphipyrinae, 175
Anacamptodes humaria humaria, 177

aberrations, 123
abnormalities, 13

Abagrotis hennei, 73 Anaea, 120
Acontinae, 176 Anartia, 37
Acraea bonasia, 154 A. fatima, 58

Acronicta arioch, 196 A. jatrophe, 59
Actias luna, 165, 262 Ancylis, 23
adaptation, 77, 85 Ancyloxypha numitor, 157
Adelpha, 121 Anderson, William A., 143
Aeropetes tulbaghia, 70 Angevine, Mark W., 29
Aganisthos, 119 Annaphila hennei, 73
aggregative behavior, 57 Antheraea, 207
Agliinae, 207 A. polyphemus polyphemus, 147, 262
Agroperina lutosa, 175 A. cooperii, 264
Agrotis ipsilon, 199 Anthocharis midea, 157
Alopex lagopus (Carnivora), 100 Apamea inebriata, 175
Amblyscirtes belli, 189 Apodemia, 74
A. hegon, 157 A. mormo, 259
A. samoset, 150 Archaeoattacus, 207
Amphicoma cooperii, 264 Archaeoprepona demophon centralis, 119,
Amphion nessus, 263 120

272

Arctiidae, 174, 196
_ Argema, 207
Argynnis zerene, 137
A. monticola, 138
Aricia agestis, 103
A. artaxerxes, 103
Arsenurinae, 208
Artogeia, 77
A. adalwinda thomsoni, 103
A. dulcinea, 81, 104
A. ergane, 96
A. flavescens, 104
. japonica, 81
. krueperi, 83
. marginalis, 80
melete, 79
m. melete, 81
napi adalwinda, 80
. bicolorata, 93
. britannica, 81
. bryoniae, 79
. bryoniaemelete, 77
. dubiosa, 82
. hulda, 82, 104
. macdunnoughii, 85
. marginalis, 96
. meridionalis, 02
. napi, 85
. neobryoniae, 81
. oleracea, 81
. passosi, 104
. pseudoropae, 82
. septentrionalis, 103
. venosa, 82
. narina, 104
. rapae, 84, 95
. virginiensis, 81
Asilidae (Diptera), 73
Asterocampa celtis, 158
A. c. antonia, 58
A. c. clyton, 158
Atalopedes campestris, 157
Athesis, 1
Atrytone delaware, 157
Attacus, 207
Atticidae, 207
Automeris, 208
A. 10, 196, 262
Baker, James H., the collection, 36
Battus belus varus, 56
B. philenor, 56, 157
B. polydamas, 56
Bellura melanopyga, 175
Berberis, 233
Biblis, 12]
biennialism, 167

>>> DSEE DEED DSS DD DED DEED DDD
SSS SSSSQSSSSTSPSPSPSSS

JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

Biston betularia, 60

B. b. carbonaria, 60

B. b. insularia, 60

B. b. “typical,” 60

B. cognataria, 148
Blanchard, A., 179, 209
Boisduval, Jean, 137
Boloria bellona, 158

B. chariclea, 167

B. distincta, 167

B. polaris stellata, 167

B. selene, 143
Bombycoidea, 208
Bombyx mori, 208
Bombus (Hymenoptera), 259

book reviews, 55, 69, 123, 206, 266, 268,

DD
Borbo fatuellus, 71
Borocera, 208
Bowden, S. R., 77
Bradley, Brian P., 162
Brenthis selene marilandica, 143
B. s. myrina, 143
Brother Apolinar Maria, 65
Brown, Richard L., 21
Brussard, Peter F., 29
Bubulcus ibis (Aves) 198
Bunaeini, 208
Caenurgina crassiuscula, 199
C. erechtea, 199
Calephelis, 218 |
C. muticum, 189, 190

Callophrys augustinus augustinus, 248

. comstocki, 73
. eryphon eryphon, 248, 252
_ henrici, 157
. niphon, 248
_n. Clarki, 248, 252
. polios obscurus, 248
Callopistria floridensis, 175
Callosamia, 207, 232
C. angulifera, 236
C. promethea, 232, 263
C. securifera, 237
Calycopis cecrops, 157
cannibalism, 129
carbonaria, 60
Carterocephalus palaemon, 150
Catocala, 133
. amica, 267
. gracilis, 176
. herodias, 176
. lacrymosa, 267
. micronympha, 176
. paleogama, 176, 266

QAAAAMAY

Bae: a ea

VOLUME 33, NUMBER 4 ANS

Catocalinae, 176 Ctenuchidae, 73
Celestrina argiolus pseudargiolus, 150, Cuculliinae, 129, 175
158 Cyllopsis gemma, 158

C. ebenina, 189 Cynthia, 268
Ceratomia catalpae, 263 Cyrtogone, 208
Cercophana, 208 Danaidae, 156
Cercophanidae, 208 Danaus plexippus, 158, 230
Chaetaglaea sericea, 129 Darapsa pholus, 263

C. tremula, 129 Dasycera, 139
Charaxes zoolina zoolina, 70 Deidamia inscripta, 263
Chemsak, John A., 261 Despina, 139
Chimoptesis, 23 development, 11
Chlorantha, 24 Diacrisia virginica, 196
Chlosyne nycteis, 152, 158 differential growth, 162
Choreutidae, 142 Diptera, 73, 121
Chrysopa slossonae, 243 Dolba hyloeus, 263
Citheronia regalis, 162, 166, 263 Doyle, Joseph F., III, 20

C. sepulcralis, 165 Drasteria, 136
Clarke, J. F. Gates, 36, 139 dry season, 58
Clarke, Sir Cyril A., 54, 60 Drosophila (Diptera), 99, 107
Clench, Harry K., 216, 268 Dynamine dyonis, 20
Clossiana, 90 Eacles imperialis, 165, 262

C. euphrosyne, 90 ecdysis, 57

C. selene, 90 Ecpantheria scribonia, 196
cochlid, 196 Emmel, John F., 73
Coea acheronta, 112 Emmel, Thomas C., 73
Coenonympha tullia, 78, 105 Enargia, 129
Coleoptera, 264 Endromis, 208
Colias, 85, 198 Ennominae, 177

C. croceus, 93 Epargyreus clarus, 157, 230

C. eurytheme, 47, 93, 157, 197, 199,258 Epia, 208

C. philodice, 157, 197, 199 Epiblema abruptana, 21, 184
Colobura dirce, 112 E. deflexana, 21, 184
Coloburini, 112 E. discretivana, 21
Coloradia pandora lindseyi, 167 E. exacerbatricana, 21, 27
Colotis ione, 70 E. grossbecki, 21, 184
Cooper, James Graham, 264 E. insidiosana, 21
Copaxa, 207 E. luctuosana, 21, 179, 182, 184
Cophura hennei, 73 E. minutiana, 179, 183
Copicucullia mcdunnoughi, 73 E. numerosana, 21, 184
Copiopteryx, 207 E. praesumptiosa, 21, 184
correction, 137 E. separationis, 21
Coryphista meadii atlantica, 177 E. strenuana, 179
Coscinocera, 207 Epiglaea apiata, 129
Cosmia, 129 E. decliva, 129
Cossidae, 146 Epimecis virginaria, 177
courtship behavior, 42 Epinotia dietziana, 21
Covell, Charles V., Jr., 153, 189 E. hopkinsonana, 24
Cressonia juglandis, 263 E. subviridis, 24
Crenis, 70 Epiphora, 207

C. boisduvali, 70 Epizeuxis concisa, 176
Cross, Winifred, 50 Erebia, 102
Cryphia villificans, 176 E. claudina, 167
crypts, 239 E. ligea, 167
Cryptocala acadiensis, 37 Erinaea chlorantha, 21

Cryptolechia phoenissa, 142 Erora laeta, 149, 189, 230

274

Erynnis, 218, 226

E. baptisiae, 190, 258

E. brizo, 157

E. horatius, 157

E. icelus, 150, 157

E. juvenalis, 157

E. lucilius, 189, 190
Estigmene acrea, 196, 197

E. rimosalis, 196
Euchloe, 94
Eucirroedia pampina, 129
Eucosma antaxia, 179
. argyrocyma, 204
. atascosana, 210
. cataclystiana, 209
. diabolana, 210
. graziella, 212
guttulana, 210
. hennei, 73
minutana, 179
. mirosignata, 215
. robinsonana, 214
. sandiego, 213
Eucosmini, 21, 209
Eudaemonia, 207
Eudia, 207
Eumorpha pandorus, 263
Eunica, 70
Euphydryas, 74, 218
. anicia, 170
. chalcedona, 199
.c. guino form “hennei”
.c. macglashanii, 200
. editha, 199
. phaeton, 203, 230
Euphyes dion dion, 189

E. dukesi, 189

E. vestris metacomet, 157
Eupithecia slossonata, 177
Euploea core amymone, 50
Euproserpinus euterpe, 73
Eupsilia cirripalea, 129

E. morrisoni, 129, 133

E. sidus, 129

E. transversa, 130

E. vinulenta, 129
Euptoieta claudia, 158, 230
Euptychia hermes sosybius, 158
Eurema, 85

E. lisa, 47, 157

E. nicippe, 157
Euristrymon ontario, 151
Eustera, 207
Everes comyntas, 157, 197
Evergestis rimosalis, 196
Ferguson, Douglas C., 192
Ferris, Clifford D., 123

Hahah

JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

fertility, 11

foodplants, 20, 162, 170, 199
Freeman, H. A., 124
Freytag, Paul H., 153
Gelechiidae, 140

gene frequency, 29

Geometridae, 60, 73, 148, 177, 199, 239

Gibson, Loran D., 189
Gill, Alison, 50
Givira anna, 146
Glenoides texonaria, 177
Glyphipterygidae, 142
Glyphodes pyloalis, 196
Godfrey, George L., 198
Gonepteryx, 85
Gononeta, 208
Graellsia, 207
Graphiphora tenuicula, 175
Graphium marcellus, 157
Grapholitha vucciniana, 25
Griselda myrtillana, 21

G. radicana, 24

G. stagnana, 21

G. vacciniana, 25
Haematopis grataria, 198
Hadeninae, 175
Hamadryas, 114

H. februa, 201
Harkenclenus titus mopsus, 157
Harvey, Donald J., 151
Heliconiinae, 67
Hemaris diffinis, 256

H. gracilis, 256

H. thysbe, 256
Hemerocampa leucostigma, 196
Hemileuca, 208
Hemileucinae, 208
Henderson, Richard A., 189
Henne, Christopher, 72
Hepialidae, 192
Hepialus fusconebulosa, 192
. gracilis, 192
. humuli, 192
hyperboreus, 192
. lupulinus, 192
. mathewi, 192
. meglashani, 192
. mustelinus, 192
. pulcher, 192
. roseicoput, 192
. sciophanes, 192
Herminiinae, 176
Hesperia, 218

H. juba, 258

H. leonardus, 230

H. manitoba borealis, 167

eepsspisedsopicspicepicepcsiisopise

Hesperiidae, 71, 124, 156, 167, 258, 259

VOLUME 33, NUMBER 4

Heterocampa varia, 176
heterozygosity, 33
Historis odius, 112, 201
Homoglaea hircina, 129
Holomelina opella, 174
holotype designation, 135
Hyalophora, 207

H. cecropia, 232, 263
hybridization, 6
Hydriomena transfigurata, 177
Hylephila phyleus, 230
Hyles lineata, 263
Hymenoptera, 121, 154, 264
Hypeninae, 176
Hypenodes fractilinea, 176
Hyperskeles choreutidea, 139, 140
Hyphantria cunea, 196
Hypolimnas bolina, 50
Hypomecis gnopharia, 177
Hypothyris daeta, 69

H. euclea leucania, 68
Imbrasia, 207
Inguromorpha basalis, 146
Incisalia, 248
Irwin, Roderick R., 268
Isia isabella, 196
Ithomiinae, 1, 67
Jodia rufago, 130
Khalaf, Kamel T., 196
Korscheltellus lupulinus, 192
Kundrya finiti mana, 21
Lamas, Gerardo, 1, 65
Lapara coniferarum, 263
Larentiinae, 177
Lasiocampa quercus callunae, 167
lasiocampid, 196
Lecithoceridae, 206
Leeuw, Irwin, 204, 267
Leptidea synopsis, 47
Lethe anthedon, 191

L. appalachia, 29, 191

L. creola, 191

L. eurydice, 29

L. portlandia, 35, 189, 191

L. p. missarkae, 189, 191
Leptotes marina, 189, 190

L. m. “burdicki,” 73
Leucania, 198

L. linda, 175
Libythaena bachmanii, 156, 158
Libytheidae, 156
life cycle, 112
life history, 254
Limenitis arthemis astyanax, 158
liperid, 196
Lithophane, 129

L. baileyi, 130

275

. bethunei, 129
. grotei, 129
. hemina, 129
innominata, 129
. patefacta, 129, 133
. petulca, 129
. querquera, 129, 133
. semiusta, 129, 133
. signosa, 129
. tepida atincta, 130
Lithophanini, 129
Lomographa glomeraria, 177
Luperina passer, 175
Lycaena hyllas, 157
L. phlaeas americana, 157
Lycaeides melissa annetta, 265
L. m. melissa, 265
Lycaenidae, 73, 115, 149, 151, 156, 248,
265
Lymantria dispar, 50, 133
Lytta cooperi, 264
macrolepidoptera, 174, 203
Malacosoma, 130
M. disstria, 196
malaise trap, 153
malfunction, 57
Manduca quinquemaculata, 263
M. sexta, 263
Maniola jurtina, 78, 105
Marpesia berania, 59
Masters, John H., 135, 137, 167, 199, 265
Mattea, 139, 142
McCabe, Timothy L., 37
McInnis, Michael L., 189
McKillop, William Brian, 147
Mechanitini, 1
Mechanitis, 68
Megistanis, 119
Megisto cymela, 158
Melanargia galathea, 92
melanic males, 148
Melinaea mnasias, 5
Milipotis idomita, 136
M. cooperi, 264
Melitea quino, 264
Mellicta athalia athalia, 197
M. a. celadussa, 97
Meneris, 70
Mermis (Nematoda), 112
Mestra, 121
Metalectra richardsi, 176
Metaxaglaea, 129, 175
Methona, 1
microlepidoptera, 206
migratory flight, 260
Miller, Jaqueline Y., 55
Miller, Lee D., 70

SUS SIS SSeS Ss

276

Miller, William E., 204
morph systems, 79
Morphinae, 67
Morpho peleides, 202
Muller, Joseph, 174
Muyshondt, Alberto, 112
Muyshondt, Alberto, Jr., 112
Nastra lherminier, 157, 230
Nathalis iole, 222
natural history, 112
Neck, Raymond W., 57
Neil, Kenneth, 245
Nemoria obliqua hennei, 73
Neophasia menopia, 259
Nephila madagascariensis, 208
Neuroptera, 243
new combination, 204
new species, 179, 192, 209
new subspecies, 245
new status, 179
Neophasia menapia, 259
Noctua chardinyi, 37
Noctuidae, 37, 73, 129, 136, 175, 196, 199,
245, 266
nomenclatural changes, 21
Norma, 21
Notocelia culminana, 21, 27
N. illotana, 21, 27
N. purparissatana, 21, 27
N. suffusana, 21, 27
N. trimaculana, 21, 27
Notodontidae, 176, 196
Nudaurelia, 207
Numerosana, 27
Nymphalidae, 1, 6, 20, 57, 58, 73, 112, 137,
143, 156, 167, 170, 199, 200, 201, 203,
244, 259, 265
Nymphalinae, 58, 67, 202
Nymphalis, 90
N. antiopa, 90
N. californica, 200
N. milberti, 200
N. vau-album, 150, 200
obituaries, 72
occurrences, 143, 149
Ochlodes sylvanoides, 259
Ochropleura plecta, 41
Ocinara, 208
Oecophoridae, 139
Oeneis jutta, 167
O. macounii, 107
O. nevadensis, 167
Orgyia leueostigma, 245
O. |. intermedia, 247
O. 1. plagiata, 245
O. 1. sablensis, 245

JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

Olethreutinae, 21, 204
Oligia minuscula, 175
Oliver, Charles G., 6, 244
Olyra, 4
Oosting, Daniel P., 149, 151
Ophiderinae, 176
Ornithopetera priamus, 57
overwintering, 166
oviposition, 56, 203
Paedisca constrictana, 187
Paititia neglecta, |

P. exaecatus, 263
Paonias astylus, 263
Papilio acheronta, 119

P. aristodemus ponceanus, 257
P. cadmus, 120

P. cinyras, 135

P. c. brasiliensis, 136
P. c. cinyras, 135

P. c. ridens, 135

P. cresphontes, 257
P. c. cresphontes, 257
P. dardanus, 50

P. d. hippocoon, 54
P. glaucus, 50, 151

P. g. canadensis, 150
P. machaon, 19, 50

P. odius, 119

P. orion, 119

P. thoas, 135

2

. troilus, 157
Papilionidae, 56, 66, 135, 156
Papilionoidea, 71
Parahypenodes quadralis, 176
Paratrea plebeja, 263
Passer domesticus (Aves), 95
Penstemonia hennei, 73
Petrova, 204

P. argyrocyma, 204

P. khasiensis, 204
Phaneta indagatriciana, 212

P. mayelisana, 209

P. verecundana, 209
phenotype, 11
Philgalia denticulata, 177

P. strigataria, 177
Philosamia, 207
Philotes enoptes dammersi, 73
Phoebis, 59, 114
Pholisora catullus, 157
Phyciodes batesii, 6

P. campestris, 7

P. tharos, 6, 153, 158, 244
Pieridae, 42, 59, 66, 77, 156, 197, 199, 258,

259

Pieris, 77

P. brassicae, 84, 95

VOLUME 33, NUMBER 4

. bryoniae, 94

. napi, 94, 258

n. marginalis, 85

n. microstriata, 258
n. oleracea, 85

n. venosa, 85

. protodice, 42

BMa~ MaMa MaMa MacMaclac

Pinara, 208
Plathypena scabra, 198
Platt, Austin P., 28, 162
Platyperigea multifera, 175

RP extinca, 175
pleiotrophy systems, 90
Plutella xylostella, 196
Poanes hobomok, 157

P. viator, 189

P. v. zizaniae, 189

P. zabulon, 157, 189
Polites coras, 152, 157

P. origenes, 157

P. themistocles, 157
Polydorus aristolochiae, 57
Polygonia, 226

P. comma, 150, 158, 230

P. faunus, 150

P. interrogationis, 158

P. zephyrus, 259
polymorphism, 77
polyphagous, 131
Polythrix, 124
. alciphron, 124
. asine, 124
. caunus, 124
. guatemalaensis, 124
. lindora, 128
. mexicanus, 124
. octomaculata, 124
. procerus, 124
Polythysana, 208
Pompeius verna, 157
Pontia collidice, 93
populations, 153, 266
population structure, 29
Precis coenia, 158
predatory behavior, 129
Prepora, 119, 120, 201

P. omphale octavia, 120
Preston, William B., 147
Proannaphia, 73
Pseudaletia unipuncta, 198
_Pseudoboarmia, 177
Pycina zelis, 112, 120
Pyralidae, 73, 196, 199
Pyreferra ceromatica, 133

P. citrombra, 129

me} Ine} Is} Ie} Ine} Ins} Ins) Ine}

. rapae, 46, 87, 157, 197, 218
. virginiensis, 19, 87, 94, 150

P. hesperidago, 129

P. pettiti, 129
Pyrgus communis, 157
Quinta cannae, 121
Racheospila hennei, 73
range extension, 20
regional lists, 216
Reist, James D., 248
Rhopobota, 21

R. dietziana, 21
. eclipticodes, 23
. finitimana, 21
. fractifasciana, 24
. microrrhyncha, 23
. naevana, 21
. unipunctana, 21
. ustomaculana, 23

DUD DnaD

Rhynchagrotis giluipennis, 37

Riodinidae, 156, 259
Rothschildia, 207
Rutkowski, Frank, 258
Rutkowski, Ronald L., 42
Sagana, 207

Salassinae, 208

Sallya, 70

Samia, 207

Saturnia, 207

277

Saturniidae, 147, 162, 166, 196, 207, 232

Satyridae, 102, 156, 167
Satyrinae, 66

Satyrium acadica acadica, 204
S. calanus falacer, 151, 157

S. caryaevorus, 151
Schizura unicornis, 196

Schweitzer, Dale F., 129, 136, 146

selective neutrality, 85
Sericaglaea signata, 130
Sericoris myrtillana, 25
Sesiidae, 73

sex chromatin, 50

sex ratio, 11

Shapiro, Arthur M., 197, 200, 258, 260

Sibine stimulea, 196
sibling species, 29
Sicya macularia, 177
Siderone, 120
Simmons, Robert S., 143
Siproeta, 58

S. steneles, 59
Smerinthus jamaicensis, 263
“Smith” body, 52
Smyrna blomfildia, 112

S. karwinski, 59, 112
Sonia constrictana, 179, 185

S. paraplesiana, 179, 184, 185

Speyeria, 137, 218
S. coranis hennei, 73

278 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

S. cybele, 138, 152 Tortricidae, 21, 73, 179, 204, 209

S. idalia, 265

S. zerene, 138, 158
Sphecodina abbottii, 263
Sphingidae, 254
Sphinx gordius, 263
Spodoptera eridania, 196

S. latifascia, 196

S. ornithogalli, 199

S. stimulea, 196
Stamp, Nancy E., 203
Sternberg, James G., 232
Sthenodis argenteomaculatus, 195

S. auratus, 193, 195

S. quadriguttatus, 195
Strymon melinus, 157
subspecific variation, 77
Synadia, 208
Synchiora aerata, 239

S. frondaria, 239

S. liquoraria liquoraria, 239
Syntherata, 207
Tabanidae (Diptera), 154
tachinid (Diptera), 112
Tacparia atropunctata, 177
Tarachidia semiflava, 176
Taragamma, 208
Tenebrio (Coleoptera), 130
Teras verditer, 21
Thecla syncellus, 55
Theophila, 201
Therina, 208
Thorybes, 218

T. bathyllus, 218

T. pylades, 218
Thymelicus lineola, 157
Thyridia psidii, 1
Tithoreini, 4
Toliver, Michael E., 232
Tortix naevana, 21

T. unipunctana, 21

Treat, Asher E., 148
Treiber, Miklos, 239
Tribolium spp. (Coleoptera), 169
Trichoplusia ni, 121
Trigona, 202
type locality, 137
Tyriomorpha, 139, 142
ultraviolet light trap, 261
Urania fulgens, 260
Uranidae, 260
Vanesiini, 112, 247
Vanessa, 230, 268

V. atalanta, 158, 230

V. cardui, 123

V. elymi, 123

V. varini, 123

V. virginiensis, 158, 259
viability, 11
voltinism, 13
Wagner, Warren H., Jr., 151
Waldbauer, Gilbert P., 232
Wallengrenia egeremet, 157

W. etho egeremet, 152
weather, 68
wheat germ diet, 196
White, Raymond R., 170
Williams, Benjamin D., 254
Williams, Thomas F., 162
Worth, C. Brooke, 162, 166, 264
Wright, William B., 266
Xanthocleis ino, 1
Xanthopastis timais, 196
Xylena currimacula, 129
Xylomoia chagnoni, 50, 175
Young, Allen M., 56, 68, 201
yponomeutid, 196
Zale lunata, 196

Z. metata, 176

VOLUME 33, NUMBER 4

PLANT INDEX FOR VOLUME 33

Abies balsamea, 150
Acanthaceae, 58
Achillea millefolium 37
Acer nigrum, 65

A. rubrum, 151

A. saccharum, 149, 151

A. spicatum, 150
Ageratum, 240
Agrostis idahoensis, 260
Allium, 200
Ambrosia, 240

A. artes misiifolia, 181

A. alnifolia, 28
Amelanchier laevis, 37
Amorpha fruticosa, 266
Amyris elemifera, 257
Anacardiaceae, 165
Apocynum androsaemifolium, 37
Aquifoliaceae, 24
Aquilegia canadensis, 190
Arctostaphylos uva-ursi, 266
Aristolochia constricta, 57

A. ringens, 57

A. serpentaria, 151
Aristolochiaceae, 56
Artemisia californica, 240
Asiminia tribola, 151
Aster, 240

A. simplex, 7

A. laevis, 7

A. macrophyllus, 150

A. undulatus, 7
Asteraceae, 239
Baptisia tinctoria, 266
Barbarea vulgaris, 150
Barberry (Berberis) 177, 233
Besseya alpina, 170
Betula, 177
Bidens, 240

B. aristosa, 190
Borrichia fructens, 27
Brassica, 198
Carex, 30
Carya spp., 162, 176

C. tomentosa, 165

C. orata, 151

€. ovalis, 151
Castilleja lapidicola, 200

C. nana, 200

C. pilosa, 199
Caulophyllum thalictroides, 151
Ceanothus, 177

C. pubescens, 266
Cecropia mexicana, 112

C. peltata, 112

Centaurea, 198
Chelone glabra, 203
Chrysanthemum, 240
Chrysothammus nauseosus, 259
Citrus paradise, 257
Clitoria sp., 59
Colobura dirce, 112
Compositae, 27, 90
Conifer, 24
Convolvulaceae, 57
Coreopsis, 240
Coronella varia, 258
Corylus cornuta, 150°
Crataegus, 151
Cucurbitaceae, 57
Cystopteris protrusa, 151
Dentaria diphylla, 19
Diospyros virginiana, 162
Diplacus, 200
Dipsaceae, 25
Ebenaceae, 165
Embauba, 112
Ericaceae, 24
Erigeron canadensis, 240
Eupatorium, 240
Fagus grandifolia, 149
Ferns, 175
Fraxinus, 162

F. americana, 149

F. nigra, 150
Glycine max, 198
Gossypium, spp., 162
Hamamelidaceae, 165
Hamamelis virginiana, 133
Hepatica acutiloba, 151
Hypericum perforatum, 37
Ilex, 24
Ilex opaca, 175
Iris versicolor, 247
Juglandaceae, 163, 165
Juglans, 176

J. cinerea, 162

J. nigra, 162
Juniperus sibirica, 266
Laportea canadenis, 151
Leguminosae, 59, 90, 258
Liatris, 239, 241

L. graminifolia, 240

L. spicata var. resinosa, 240
Lindera benzoin, 151
Liquidambar styraciflua, 162
Macglashanii, 200
Medicago sativa, 198
Melilotus alba, 151

M. officinalis, 151

280 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

Melothria guadelupensis, 57
Mimulus, 200
Monardella, 201
Moraceae, 112
Myrica pensylvanica, 246
Nyssa sylvatica, 162
Onoclea sensibilis, 150
Opuntia polyacantha, 266
Orthocarpus, 200
Ostrya virginiana, 151
Penstemon brevifloris, 200

P. lemmonii, 200
Phalaris arundinacea, 175
Phryna leptostachya, 151
Pinus echinata, 147

P. banksiana, 248

P. contorta, 249

P. c. latifolia, 250

P. rigida, 147

P. virginiana, 176
Plantago major, 174
Platanus occidentalis, 130, 151, 162
Podophyllum peltatum, 151
Polygonaceae, 57
Potentilla fruticosa, 266
Populus deltoides, 151

P. tremuloides, 150, 248
Prunus pensylvanica, 149

P. virginiana, 37

P. serotina, 233

P. pumila, 266
Pteridium aquilinum, 37
Pulchea odorata, 240
Purpurea, 131
Pyrus, 131
Quercus, 152, 162, 176

Q. rubra, 150, 151, 176

QO. velutina, 266
Rhododendron, 195
Rhus spp., 162

R. capallina, 162
Rosa, 247, 248
Rubus idaeus, 37

R. parviflorus, 150

Rudbeckia hirta, 240
Rumex, 37

R. occidentalis, 175
Ruta graveolens, 257
Sablensis, 247
Sagittaria latifolia, 37
Sanguinaria canadensis, 151
Salix, 162, 204, 266
Sambucus canadensis, 37
Sassafras, 177
Scabiosa columbaria, 25
Scrophularia, 200

Scrophulariaceae, 170, 199, 203

Sisymbrium iris, 42
Solanaceae, 58
Solanum rugosum, 68
Solidago, 239 |

S. canadensis, 240

S. nemoralis, 240

S. pinetorum, 240
Spirea, 177

S. latifolia, 37
Sterculiaceae, 201
Syringa vulgaris, 162
Taraxacum officinale, 174, 259
Theobroma cacao, 201
Tilia americana, 149
Tragia ramosa, 20
Trifolium hybridum, 189
Trillium cernuum, 150
tulip tree, 177
Ulmus americana, 149
Urera caracasana, 112
Urticaceae, 112
Vaccinium, 176, 247

V. corymbosum, 195, 255

V. macrocarpon, 24

V. myrtillana, 25

V. myrtilloides, 7

V. vacillans, 254
Viola pedatifida, 265
yellow water lily, 175

Xanthoxylum americanum, 151

Date of Issue (Vol. 33, No. 4): 8 April 1980

EDITORIAL STAFF OF THE JOURNAL
AUSTIN P. PLATT, Editor
Department of Biological Sciences

University of Maryland Baltimore County, 5401 Wilkens Avenue
Catonsville, Maryland 21228 U.S.A.

FRANCES S. CHEW, Managing Editor
Department of Biology
Tufts University
Medford, Massachusetts 02155 USA

DOUGLAS C. FERGUSON, Associate Editor THEODORE D. SARGEN’, Associate Editor

NOTICE TO CONTRIBUTORS

Contributions to the Journal may deal with any aspect of the collection and study of
Lepidoptera. Contributors should prepare manuscripts according to the following in-
structions.

Abstract: A brief abstract should precede the text of all articles.

Text: Manuscripts should be submitted in triplicate, and must be typewritten,
entirely double-spaced, employing wide margins, on one side only of white, 8% x 11

_ inch paper. Titles should be explicit and descriptive of the article’s content,including

the family name of the subject, but must be kept as short as possible. The first mention
of a plant or animal in the text should include the full scientific name, with authors
of zoological names. Insect measurements should be given in metric units; times
should be given in terms of the 24-hour clock (e.g. 0930, not 9:30 AM). Underline only
where italics are intended. References to footnotes should be numbered consecutively,
and the footnotes typed on a separate sheet.

Literature Cited: References in the text of articles should be given as, Sheppard
(1959) or (Sheppard 1959, 1961la, 1961b) and all must be listed alphabetically under
the heading LITERATURE CITED, in the following format:

SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd. ed. Hutchinson, London.
209 p. é

196la. Some contributions to population genetics resulting from the study of
_ the Lepidoptera. Adv. Genet. 10: 165-216.

In the case of general notes, references should be given in the text as, Sheppard (1961,
Adv. Genet. 10: 165-216) or (Sheppard 1961, Sym. R. Entomol. Soc. London 1: 23-30).

Illustrations: All photographs and drawings should be mounted on stiff, white
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er. The author’s name, figure numbers as cited in the text, and an indication of the
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Tables: Tables should be numbered consecutively in Arabic numerals. Headings
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Correspondence: Address all matters relating to the Journal to the editor. Short
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PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044 U.S.A.

CONTENTS

FIVE NEW SPECIES OF THE TRIBE EUCOSMINI (TORTRICIDAE).
André Blanchard 030

How TO MAKE REGIONAL LISTS OF BUTTERFLIES: SOME
THOUGHTS. Harry K. Clench 02) 3

DAILY FLIGHT PERIODS OF MALE CALLOSAMIA PROMETHEA
(SATURNIIDAE). Michael E. Toliver, James G. Sternberg &
Gilbert P. Waldbauer 0 ee

COMPOSITES AS HOST PLANTS AND CRYPTS FOR SYNCHLORA
AERATA (GEOMETRIDAE). Miklos Treiber... Dee

A NEW SUBSPECIES OF ORYGIA LEUCOSTIGMA (LYMANTRIIDAE)
FROM SABLE ISLAND, NOVA SCOTIA. Kenneth Neil ________-

CALLOPHRYS NIPHON (LYCAENIDAE) IN ALBERTA WITH NOTES ON
THE IDENTIFICATION OF C. NIPHON AND C. ERYPHON. ones
Dy Feist ie I

LIFE HISTORY OBSERVATIONS ON HEMARIS GRACILIS (SPHINGI-
DAE). Benjamin D. Williams |...

GENERAL NOTES
A new method of inducing copulation in Phyciodes tharos (Nymphalidae).
Charles Gy OlWoep 0a ne ee el ao See
New Papilio cresphontes hostplant. Frank Rutkowski ____.-.-.----------_----

Erynnis baptisiae (Hesperiidae) on crown vetch (Leguminosae). Arthur
M. Shapiro’ foo ns

Does Hesperia juba (Hesperiidae) hibernate as an adult? Arthur M.
Shapiro 22 i NY

A migratory flight of Urania fulgens (Walker) in Honduras (Uraniidae).
John A. Chemsak, E. G. Linsley & Juanita M. Linsley _-.--..1.- 2

Captures of large moths by an ultraviolet light trap. C. Brooke Worth &
Joseph Maller 2.00 a ea

James Graham Cooper (1830-1902). John H. Masters ____-----------------_-

A recent record of -Speyeria idalia (Nymphalidae) from Manitoba. John
H. Masters 2222.0.0 2) a an

Aberrant specimen of Lycaeides melissa melissa (Lycaenidae). William B.
Wright -2c TCR SN a aT) er

Population outbreak of Catocala palaeogama (Noctuidae). Irwin Leeuw _

Notes on the biology of Battus philenor (Papilionidae) in Centre County,
Pennsylvania. Frank D. Fee & Richard Boscoe __---.---------+1----2-2

BooK REVIEWS

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